new and interesting Cantharellus from tropical Africabakasbl.org/news/doc/194.pdf · 485 / DS 06.033 Malaysia PC0084739 – KF294740 C.eucalyptorum Buyck & V. Hofstetter 2015 59

Cryptogamie, Mycologie, 2016, 37 (3): 283-327© 2016 Adac. Tous droits réservés

doi/10.7872/crym/v37.iss3.2016.283

new and interesting Cantharellus from tropical Africa

André DE KESELa*, Mario AMALFI a, Bill KASONGO WA NGOY b,Nourou Soulemane YOROU c, Olivier RASPÉ a, d,

Jérôme DEGREEF a, d & Bart BUYCK e

aBotanic Garden Meise, Nieuwelaan 38, 1860 Meise, Belgium,email: [email protected]

bFaculty of Agronomy, University of Lubumbashi, 1825 Kasapa Road,Lubumbashi, DR Congo

cFaculty of Agronomy, University of Parakou,Dept. of Natural Resources Management, BP 123, Parakou, Bénin

dFédération Wallonie-Bruxelles, Service Général de l’Enseignement universitaireet de la Recherche scientifique, rue A. Lavallée 1, 1080 Bruxelles, Belgium

eMuséum national d’Histoire naturelle, Département Systématique et Évolution,CP 39, ISYEB, UMR 7205 CNRS MNHN UPMC EPHE,

12 rue Buffon, F-75005 Paris, France, email: [email protected]

Abstract – This paper deals with some of the larger, more or less yellowish or orangeCantharellus species from the tropical African woodlands and rain forests. Four new specieswith clamp connections are described: Cantharellus guineensis, C. mikemboensis,C. pseudomiomboensis and C. stramineus. The new taxa show moderate to strong resemblanceto either Cantharellus rufopunctatus or C. miomboensis. A two-locus phylogeny, based onpart of the protein coding genes rpb2 and tef-1, resolved them as a highly supported cladewithin Cantharellus subgenus Rubrinus, a subgenus still exclusively composed of tropicalAfrican species. This monophyletic clade is here described as a new section within subg.Rubrinus. As the subgenus was previously defined as being composed of chanterelles lackingclamp connections, the definition of the subgenus is here amended. Illustrations and newrecords are also presented for Cantharellus afrocibarius, C. defibulatus, C. miomboensis,C. rufopunctatus and C. sublaevis. This paper provides first sequences for C. defibulatus,C.rufopunctatus and C. sublaevis, all of which are here epitypified, as well as new sequencesfor more than a dozen other Cantharellus. Cantharellus cibarius var. latifolius is considereda synonym of C. afrocibarius. An identification key to all mainland African Cantharellus isproposed.

Cantharellales / Cantharellus rufopunctatus / tef1-alpha / identification key / miombo /phylogeny / rain forest / rPB2 / taxonomy

* Corresponding author: [email protected]

284 A. De Kesel et al.

IntrodUCtIon

Recent studies on Cantharellus Adans.:Fr. from tropical Africa (Zambia,Tanzania) and Madagascar describe a substantial number of new taxa from woodlandvegetation (Ariyawansa et al. 2015; Buyck et al. 2000, 2012, 2013, 2014, 2015;Buyck 1994, 2014, Buyck & Randrianjohany, 2013; Eyssartier & Buyck 1999a,b;Eyssartier et al. 2002; Liu et al. 2015; Tibuhwa et al. 2008, 2012). A worldwidephylogeny was provided to accommodate most of these new African taxa (Buycket al. 2014) and demonstrated that the bulk of the African chanterelles belong to twowell-supported monophyletic clades that are entirely composed of species lackingclamp connections. One of the earliest described species among these clamplesschanterelles, Cantharellus rufopunctatus (Beeli) Heinem., is probably one of themost cited but least understood taxa. Originally described from the Guineo-Congolianrainforest (Beeli 1928, Heinemann 1958), we will show here that this name has laterbeen systematically misapplied to taxa from woodland vegetation in both theZambezian and Sudanian ecozones of tropical Africa (Heinemann 1966, Buyck1994, Buyck et al. 2000, Eyssartier 2001, De Kesel et al. 2002, De Kesel 2004, Eyiet al. 2011, Sharp 2011, Härkönen et al. 2015). Buyck et al. (2013) were the first toresolve part of the puzzle by describing Cantharellus miomboensis Buyck &V. Hofstetter, based on Tanzanian and Zambian material that had previously beenidentified as C. rufopunctatus (Buyck et al., 2000). Nevertheless, even today theboundaries around C. rufopunctatus remain unclear and confusingly wide. The mainreasons for this are (1) the fact that this species has never been recollected from itsoriginal habitat and (2) the impossibility to obtain sequences from the type collection.

During recent collecting trips to both the Guineo-Congolian rainforestand the miombo woodlands of the Democratic Republic of Congo (DR Congo),special emphasis was put on recollecting C. rufopunctatus, C. miomboensis andmorphologically similar species. The objective of this paper was to designate asuitable epitype for C. rufopunctatus, and to name and identify similar taxa usingmorphological and molecular data.

MATERIAL AND METHODS

Collecting and morphology

Specimens were collected in the field following the protocol outlined inDe Kesel (2004) and Eyi Ndong et al. (2011). The macroscopic descriptions arebased on notes and photographs taken from fresh basidiomata. Colour codes followKornerup & Wanscher (1978). Microscopic structures were revived in 5% potassiumhydroxide (KOH) and examined in Congo-red ammonia solution using an OlympusBX-51 compound microscope. Basidiospores, basidia and pileal elements wererandomly selected from the specimens and measured using a digital camera andAnalySIS Five imaging software (Soft Imaging System GmbH). Mean values(underlined) ± 1.96 × standard deviations, and extreme values (between brackets)are given for all microstructures and derived parameters (Q = ength/width ratios).

All descriptions are based on the type material. Deviations observed inspecimens other than the type are discussed in the comment sections.

New and interesting Cantharellus from tropical Africa 285

All microscopic features were drawn by hand, using a drawing tube. Typespecimens and other collections are deposited at BR (Herbarium Meise, Belgium).

DNA extraction, amplification and sequencing

The genomic DNA of 53 Cantharellus species (Table 1) was isolated fromCTAB-preserved tissues or dry specimens using a CTAB isolation procedure adaptedfrom Doyle & Doyle (1990).

For transcription elongation factor 1-alpha (tef-1), a fragment locatedbetween exons 4 and 8 (Wendland & Kothe 1997) was amplified using the primerpair 983F and 2218R (Rehner & Buckley 2005) with the exception of a fewspecimens for which these primers did not work. For those specimens we used theprimers pair tef 1 F and tef 1 R published by Morehouse et al. (2003) following theprotocol described. For the primer pair 983F and 2218R a touchdown PCR was usedwith an initial annealing temperature of 60°C following Rehner & Buckley (2005).tef-1 was amplified in 50-μL reactions mix containing 1× polymerase buffer, 200 μmof each dNTP, 200 μg μL−1 bovine serum albumin, 0.3 μm of forward and reverseprimers, 1.25 U Taq polymerase (DreamTaq™, Thermo Fisher Scientific, St. Leon-Rot, Germany).

The region located between domains 6 and 7 of the second largest subunitof the RNA polymerase II (rpb2) (Frøslev et al. 2005, Matheny 2005) was amplifiedusing the degenerate primers bRPB2-6F and bRPB2-7.1R using the PCR programdetailed in Matheny (2005).

Partial rpb2 gene was amplified in 40-μL reactions containing 1× polymerasebuffer, 1 μm MgCl2 (2.5 μm with the MgCl2 contained in the polymerase buffer),0.75M betaine (Sigma B0300, Diegem, Belgium), 200 μm of each dNTP, 200 μg μL−1bovine serum albumin, 0.3 μm of forward and reverse primers, 1.25 U Taq polymerase(DreamTaq™), using the PCR program detailed in Matheny (2005). Successful PCRreactions resulted in a single band observed on an 0.8% agarose gel, correspondingto approximately 1200 bp. PCR products were purified by adding 1 U of ExonucleaseI and 0.5 U FastAPAlkaline Phosphatase (Thermo Scientific, St. Leon-Rot, Germany)and incubating at 37°C for 1 h, followed by inactivation at 80°C for 15 min.

Sequencing was performed by Macrogen InC. (Korea and The Netherlands)with PCR primers, for tef-1 additional sequencing was performed with the twointernal primers, 1953R and 2212R (Rehner & Buckley 2005). The sequences wereassembled in Geneious Pro v. 6.0.6 (Biomatters).

Phylogenetic analysis

One hundred and forty-six specimens representing 63 species or potentialspecies/clades were included in the phylogenetic analysis. Nucleotide sequenceswere automatically aligned with Clustal X 2.0.11 (Thompson et al. 1997) withdefault settings. Materials and sequences used in this study are listed in Table 1.

The alignment was further optimized and manually adjusted as necessary bydirect examination with the software Se-Al v. 2.0a11 (University of Oxford). Theassignment of codon positions was confirmed by translating nucleotide sequencesinto predicted amino acid sequences using MacClade 4.0 (Maddison and Maddison2000) and then compared with the annotated C. cibarius sequences AFTOL ID 607.Potential ambiguously aligned segments, especially in the four introns present in tef-1, were detected by Gblocks v0.91b (Castresana 2000; http://molevol.cmima.csic.es/


Table 1. List of species, specimens and GenBank accession numbers for sequences included inthe phylogenetic analyses. Collectors abbreviated as BB Bart Buyck, AV Annemieke Verbeken,DS Dirk Stubbe, DT Donatha Tibuhwa, GE Guillaume Eyssartier, JV Jan Vesterholt, ADK AndréDe Kesel, JD Jérôme Degreef, VDKO Omer Van de Kerckhove, KMP Kapepula Mutwale Paulin,MF Matthew Foltz, TIAN Tian Xao-Fei & ZJP Zhang Jie Ping. The newly generated sequencesare in bold

Genus / species namesVoucher extractionnr / collector nr.

Origin Herb.Acc.Nr.GenBank accession nr

tef1-α rpb2

Cantharellus addaiensis Henn. 1898AdK 3983 Kenya Br5020162751829 KX834362 KX834417AdK 4371 togo Br5020163708693 – KX834442AdK 5377 rd Congo Br5020184179731 KX834363 KX834418Jd 1093 rd Congo Br5020172953022 KX834364 KX834419267 / BB 98.057 Tanzania PC0084718 JX192976 KF294695495 / BB 98.033 Neotype Tanzania PC0084717 JX192992 KF294745

C. afrocibarius Buyck & V. Hofstetter 2012496 / BB 96.235 Holotype Zambia PC0084124 JX192993 KF294746497 / BB 96.236 Zambia PC0084125 JX192994 KF294747AdK 5389 rd Congo Br5020184182533 KX834409 KX834420AdK 5418 rd Congo Br5020184185626 KX834365 KX834421Jd 937 rd Congo Br5020169426645 KX834366 KX834422

C. albidolutescens Buyck & V. Hofstetter 2014456 / BB 08.057 Madagascar PC0084750 KF294752 KF294722457 / BB 08.070 Holotype Madagascar PC0084751 JX192982 KF294723

C. alborufescens (Malençon) Papetti & S. Alberti 1998347 / BB 07.221 USA PC0084085 GQ914939 KF294702348 / BB 07.283 USA PC0084090 GQ914945 KF294713

C. altipes Buyck & V. Hofstetter 2011318 / BB 07.019 Holotype USA PC0084085 GQ914939 KF294702344 / BB 07.162 Paratype USA PC0084090 GQ914945 KF294713

C. ambohitantelyensis Buyck & V. Hofstetter 2014475 / BB 08.336 Holotype Madagascar PC0084754 JX192989 KF294733

C. amethysteus (Quél) Quél. 1888349 / BB 07.284 Slovakia PC0084070 GQ914953 KF294716352 / BB 07.309 Slovakia PC0084071 GQ914954 KF294719

C. appalachiensis R.H. Petersen 1971342 / BB 07.123 USA PC0084075 GQ914979 KF294711

C. cerinoalbus Eyssart. & Walleyn 2009487 / AV 06.051 Isotype Malaysia PC0084743 – KF294741

C. cibarius Fr. :Fr. 1821AFTOL-ID 607 – DQ059050 DQ366285351 / BB 07.300 Slovakia PC0084077 GQ914950 KF294718479 / GE 07.025 France PC0084088 GQ914949 KF294736VdKo 1135 Belgium Br5020169491322 KX834367 KX834423




tef1-α rpb2

C. cibarius var. roseocanus Redhead, Norvell & Danell, 1997MF CC29 USA – JX030415 –

C. cinnabarinus (Schwein.) Schwein.1834312 / BB 07.001 Neotype USA PC0084094 GQ914985 KF294698326 / BB 07.053 USA PC0084093 GQ914984 KF294705

C. congolensis Beeli 1928AdK 5214 rd Congo Br5020184167660 KX834368 KX834424AdK 5215 rd Congo Br5020184166632 KX834369 KX834425

C. conspicuus Eyssart., Buyck &Verbeken 2002501 / GE 99.560 Isotype Zimbabwe PC0084809 – KF294751

C. decolorans Eyssart. & Buyck 1999469 / BB 08.278 Epitype Madagascar PC0084098 GQ914968 KF294731

C. defibulatus (Heinem.) Eyssart. & Buyck 2001AdK 5511 Epitype rd Congo Br5020184192549 KX834370 KX834426AdK 6070 rd Congo Br5020184215583 KX834371 KX834427

C. densifolius Heinem. 1958258 / BB 98.013 Tanzania PC0084126 JX193014 KF294690

C. diminutivus Corner 1969485 / DS 06.033 Malaysia PC0084739 – KF294740

C.eucalyptorum Buyck & V. Hofstetter 201559 / BB 06.148 Madagascar PC0084127 JX192965 KF29467860 / BB 06.149 Madagascar PC0084128 JX192966 KF294679

C. fistulosus Tibuhwa & Buyck 2008517 / DT 43 Isotype Tanzania PC0084738 JX192992 –

C. flavus Foltz & T.J. Volk 2013MF C067 USA – JX030416 –

C. friesii Quél. 1872481 / GE 07.077 France PC0084719 – KF294737VdKo 1165 Germany Br5020173408699 KX834408 KX881922

C. gracilis Buyck & V. Hofstetter 2012251 / BB 98.234 Holotype Tanzania PC0084737 JX192970 KF294686

C. guineensis de Kesel & Yorou sp. nov.ADK 3005 Holotype Benin Br5020129217139 KX834372 KX834428AdK 3366 Benin Br5020152165698 KX834373 KX834429

Table 1. List of species, specimens and GenBank accession numbers for sequences included inthe phylogenetic analyses. Collectors abbreviated as BB Bart Buyck, AV Annemieke Verbeken,DS Dirk Stubbe, DT Donatha Tibuhwa, GE Guillaume Eyssartier, JV Jan Vesterholt, ADK AndréDe Kesel, JD Jérôme Degreef, VDKO Omer Van de Kerckhove, KMP Kapepula Mutwale Paulin,MF Matthew Foltz, TIAN Tian Xao-Fei & ZJP Zhang Jie Ping. The newly generated sequencesare in bold (continued)




tef1-α rpb2

AdK 3489 Benin Br5020152044450 KX834374 KX834430AdK 3525 Benin Br5020152005062 KX834375 KX834431

C. heinemannianus Eyssart. & Buyck 1998491 / BB 96.307 Zambia PC0084720 – KF294743

C. hygrophorus S. C. Shao, Buyck & F. Q. Yu 2014HKAS80614 China – KJ004003

C. humidicolus Buyck & V. Hofstetter 2013493 / BB 98.036 Holotype Tansania. PC0084724 JX193005 KF294744

C. ibityensis Buyck & V. Hofstetter 2014462 / BB 08.196 Holotype Madagascar PC0084109 GQ914980 KF294727463 / BB 08.203 Paratype Madagascar PC0084722 JX192985 KF294728

C. isabellinus var. parvisporus Eyssart. & Buyck 2000256 / BB 98.020 Holotype Tanzania PC0084753 JX192972 KF294688249 / BB 98.037 Paratype Tanzania PC0084100 GQ914966 KF294685

C. lateritius (Berk.) Singer 1949320 / BB 07.025 Epitype USA PC0084103 GQ914957 KF294703330 / BB 07.058 USA PC0084105 GQ914959 KF294708

C. lewisii Buyck & V. Hofstetter 2011301 / BB 02.197 paratype USA PC0084073 GQ914961 KF294697314 / BB 07.003 holotype USA PC0084074 GQ914962 KF294700

C. luteostipitatus Buyck & V. Hofstetter 2015

464 / BB 08.210 Madagascar PC0085130 JX193008 –

C. mikemboensis de Kesel & degreef sp. nov.AdK 6039 rd Congo Br5020184208660 KX834376 KX834432AdK 6065 rd Congo Br5020184214555 KX834377 KX834433AdK 6073 rd Congo Br5020184217648 KX834378 KX834434Jd 866 rd Congo Br5020169371099 KX834379 KX834435JD 918 Holotype rd Congo Br5020169410484 KX834380 KX834436

C. minor Peck 1872313 / BB 07.002 USA PC0084747 JX192978 KF294699329 / BB 07.057 USA PC0084721 JX192979 KF294707

C. miomboensis Buyck & V. Hofstetter 2012AdK 3908 Kenya Br5020162711427 KX834381 KX834437AdK 5392 rd Congo Br5020184183561 KX834382 KX834438AdK 6006 rd Congo Br5020184197698 KX834383 KX834439





tef1-α rpb2

255 / BB 98.021 Holotype Tanzania PC0084748 JX192971 KF294687Jd 877 rd Congo Br5020169379170 KX834384 KX834440Jd 895 rd Congo Br5020169390281 KX834385 KX834441

C. nigrescens Buyck & V. Hofstetter 201519 / BB 06. 166 Madagascar PC0084979 JX192998 –66 / BB 06.176 Madagascar PC0084078 JX192967 KF29468069 / BB 06.197 Madagascar PC0084076 GQ914982 KF294683

C. nigrescens aff.AdK5441 rd Congo Br5020184187682 KX834386 –AdK5471 rd Congo Br5020184190484 KX834387 –247 / BB 98.039 Tanzania PC0084123 JX193015 –512 / BB 98.058 Tanzania PC0084776 JX192996 –

C. pallens Pilát 1959998 / BB 09.430 Italy PC0084769 KX834410 KX834463999 / BB 09.441 Italy PC0084788 KX834411 KX834464

C. paucifurcatus. Buyck & V. Hofstetter 2014474 / BB 08.320 Holotype Madagascar PC0084729 JX192988 KF294732

C. phasmatis Foltz & T.J. Volk 2013MF C057 USA – JX030417MF C074 USA – JX030418

C. platyphyllus Heinem. 1966Jd857 rd Congo Br5020169362004 KX834388 KX834444Jd919 rd Congo Br5020169411498 KX834389 KX834445Jd1107 rd Congo Br5020172967166 KX834390 –259 / BB 98.012 Tanzania PC0084108 GQ914969 KF294691262 / BB 98.126 Epitype Tanzania PC0084723 JX192975 KF294694

C. platyphyllus subsp. Bojeriensis Eyssart. & Buyck 1999458 / BB 08.158 Madagascar PC0084741 JX192983 KF294724459 / BB 08.160 Madagascar PC0084740 JX192984 KF294725

C. pseudominimus Eyssart. & Buyck 1999477 / JV 00.663 Portugal PC0084725 JX192991 KF294735

C. pseudomiomboensis de Kesel & Kasongo sp. nov.AdK 6029 rd Congo Br5020184202484 KX834391 KX834446JD 927 Holotype rd Congo Br5020169418565 KX834392 KX834447

C. quercophilus Buyck, Lewis, Eyssart & V. Hofstetter 2010636/BB 07.097 Holotype USA PC0084726 JX192981





tef1-α rpb2

C. romagnesianus Eyssart. & Buyck 1999

482 / GE 07.031 France PC0084735 – KF294738

C. rufopunctatus (Beeli) Heinem. 1958AdK 5892 rd Congo Br5020184193577 KX834393 KX834448AdK 5949 Epitype rd Congo Br5020184194604 KX834394 KX834449

C. sebosus Buyck & V. Hofstetter 2014460 / BB 08.162 Paratype Madagascar PC0084079 GQ914981 KF294726465 / BB 08.234 Holotype Madagascar PC0084736 JX192986 KF294729

C. sp. ined. (aff. splendens)499 / BB 96.199 Zambia PC0084730 – KF294749498 / BB 96.306 Zambia PC0084731 – KF294748

C. spectaculus Foltz & T.J. Volk 2013C081 USA JX030414 –

C. splendens Buyck 1994AdK 6071 rd Congo Br5020184216610 KX834395 KX834450BB 5518 Holotype Burundi Br5020059800487 – KX834443Jd 896 rd Congo Br5020169391295 KX834396 KX834451Jd 968 rd Congo Br5020169449873 KX834397 KX834452

C. stramineus de Kesel sp. nov.ADK 6051 Holotype rd Congo Br5020184210434 KX834398 KX834453Jd 1035 rd Congo Br5020172909555 KX834399 KX834454Jd 1098 rd Congo Br5020172958072 KX834400 KX834455

C. subamethysteus Eyssart. & Stubbe 2009488 / DS 06.218 Isotype Malaysia PC0084744 – KF294742

C. subcyanoxanthus Buyck, Randrianjohany & Eyssart. 2012476 / BB 00.1137 Holotype Madagascar PC0084746 JX192990 KF294734

C. subcyanoxanthus aff.257 / BB 98.014 Tanzania PC0084745 JX192973 KF294689

C. subincarnatus subsp. rubrosalmoneus Buyck & V. Hofstetter 201413 / BB 06.080 Holotype Madagascar PC0084727 JX192962 KF29467555 / BB 06.096 Paratype Madagascar PC0084755 JX192963 KF294676

C. sublaevis Buyck & Eyssart. 2014AdK 6057 Epitype rd Congo Br5020184211462 KX834401 KX834456Jd 942 rd Congo Br5020169429677 KX834402 KX834457Jd 964 rd Congo Br5020169445837 KX834403 KX834458





tef1-α rpb2

C. symoensii Heinem. 1966260 / BB 98.011 Tanzania PC0084113 GQ914970 KF294692261 / BB 98.113 Epitype Tanzania PC0084756 JX192974 KF294693KMP 2 rd Congo – KX834404 KX834459Jd 912 rd Congo Br5020169405435 KX834405 KX834460

C. tabernensis Feibelman & Cibula 1996328 / BB 07.056 USA PC0084115 GQ914974 KF294706340 / BB 07.119 USA PC0084116 GQ914976 KF294709333 / BB O7.064 USA PC0084120 GQ914975 JN993600

C. tanzanicus Buyck & V. Hofstetter 2012268 / BB 98.040 Holotype Tanzania PC0084728 JX192977 KF294696Jd 867 rd Congo Br5020169372102 KX834406 KX834461Jd 878 rd Congo Br5020169380183 KX834407 KX834462

C. tenuithrix Buyck & V. Hofstetter 2011322 / BB 07.035 Paratype USA PC0084087 GQ914946 KF294704343 / BB 07.125 Holotype USA PC0084084 GQ914947 KF294712

C. texensis Buyck & V. Hofstetter 2011317 / BB 07.018 Holotype USA PC0084097 GQ914988 KF294701341 / BB 07.120 Paratype USA PC0084096 GQ914987 KF294710

C. tomentosus Eyssart. & Buyck 2000248 / BB 98.038 Paratype Tanzania PC0084121 GQ914965 KF294684500 / BB 98.060 Holotype Tanzania PC0084732 JX192995 KF294750

C. tricolor Buyck & V. Hofstetter 201567 / BB 06.179 Madagascar PC0084129 JX192968 KF29468168 / BB 06.180 Madagascar PC0084130 JX192969 KF294682

C. tuberculosporus M. Zang 1980TIAN 351 China HKAS 55782 KM893853 –ZJP 17 China HKAS 58215 KM893854 –ZJP 117 China HKAS 58225 KM893855 –

C. variabilicolor Buyck & V. Hofstetter 201556 / BB 06.145 Madagascar PC0084111 JX193003 –57 / BB 06.146 Madagascar PC0084757 JX192964 KF29467720 / BB 06.168 Madagascar PC0124633 JX192999 –466 / BB 08.243 Madagascar PC0084733 JX192987 KF294730

Craterellus tubaeformis (Fr.) Quél.350 / BB 07.293 Slovakia PC0084122 GQ914989 KF294717



castresana/Gblocks.html, settings “allow smaller final blocks”, “allow gaps withinblocks”) and excluded from the analyses. The rpb2 or tef-1 sequences from somecollections could neither be obtained nor gathered fromGenBank and were consideredas missing data in the combined dataset. To detect the possible bias from substitutionsaturation, we tested the first, second and the third codon position of the coding regionanalyzed (rpb2 and tef-1) by using Xia’s test (Xia and Lemey 2009; Xia et al. 2003,as implemented in DAMBE (Xia and Xie 2001). Because the Iss.c is based onsimulation results, there is a problem with more than 32 species. To circumvent thisproblem, DAMBE was used to randomly sample subsets of 4, 8, 16 and 32 OTusmultiple times and perform the test for each subset to see if substitution saturationexists for these subsets of sequences. In order to confirm the results of the Xia’smethod we also plotted transitions and transversions at the first, second, and thirdcodon positions against Tamura-Nei genetic distances with the aid of the DAMBEpackage, with an asymptotic relationship indicating the presence of saturation.

The dataset was subdivided into 7 data partitions: tef-1-1st, -2nd, -3rdcodon positions, tef-1 introns, and rpb2 1st, -2nd, -3rd codon positions. Phylogeneticanalyses were performed separately for each individual and concatenated loci usingBayesian inference (BI) as implemented in MrBayes v3. 2 (Ronquist & Huelsenbeck2003, Ronquist et al. 2011) and Maximum likelihood (ML) as implemented inRAxML 7.2.7 (Stamatakis et al. 2008). Craterellus tubaeformis was used as outgroup(Buyck et al. 2013, 2014).

Models of evolution for BI were estimated using the Akaike informationcriterion (AIC) as implemented in Modeltest 3.7 (Posada and Crandall 1998). Inorder to facilitate the data partitioning by codon position for the tef-1, the fourintrons present were excised and analyzed as a distinct partition. The best-fit modelsfor each partition were implemented as partition specific models within partitionedmixed-model analyses of the combined dataset (Table 2). All parameters wereunlinked across partitions. Bayesian analyses were implemented with two independentruns, each with four simultaneous independent chains for ten million generations,starting from random trees, and keeping one tree every 1000th generation. The treewith the best likelihood value (see below ML) was used as a starting tree for theBayesian analyses. All trees sampled after convergence (average. standard deviationof split frequencies < 0.01 and confirmed using Tracer v1.4 [Rambaut and Drummond2007]) were used to reconstruct a 50% majority-rule consensus tree (BC) and tocalculate Bayesian posterior probabilities (BPP). BPP of each node was estimatedbased on the frequency at which the node was resolved among the sampled treeswith the consensus option of 50% majority-rule (Simmons et al. 2004). A probabilityof 0.95 was considered significant. Maximum likelihood (ML) searches conductedwith RAxML involved 1000 replicates under the GTRGAMMAI model, with allmodel parameters estimated by the program. In addition, 1000 bootstrap (ML BS)replicates were run with the same GTRGAMMAI model. We provided an additionalalignment partition file to force RAxML software to search for a separate evolutionmodel for each dataset. Clades with Maximum likelihood bootstrap values of 75%or greater were considered supported by the data.

To detect topological conflicts among data partitions, the nodes between themajority-rule consensus trees obtained in the ML analysis from the individual data setswere comparedwith the software compat.py (available atwww.lutzonilab.net/downloads).Paired trees were examined for conflicts only involving nodes with ML BS > 75%(Lutzoni et al. 2004, Mason-Gamer and Kellogg 1996, Reeb et al. 2004).A conflict wasassumed to be significant if two different relationships for the same set of taxa (one beingmonophyletic and the other non-monophyletic) were observed in rival trees.


rESULtS

Phylogenetic analysis

The final combined DNA sequence alignments of these two loci resultedin 1817 characters (tef1: 1030 characters, of which 626 in the exon partition and 404in the combined introns partition, rpb2: 787 characters), including gaps. Summarystatistics of sequence data and tests of substitution saturation for each dataset areprovided in Table 2. No conflict involving significantly supported nodes was foundbetween the tree topologies obtained for the individual datasets, using the 75% MLBP criterion; the datasets were therefore combined.

The test of substitution saturation (Table 2) showed that the observed indexof substitution saturation (Iss) for the tef-1and rpb2 alignments was significantlylower than the corresponding critical index substitution saturation (Iss.c), indicatingthat there was little saturation in our sequences (P < 0.001).

Twenty characters in the tef1 introns dataset were judged too ambiguous tobe aligned and excluded from the analyses. The two Bayesian runs converged tostable likelihood values after 1261000 generations. 4739 stationary trees from eachanalysis were used to compute a 50% majority rule consensus tree and to calculateposterior probabilities (PP). In the ML searches with RAxML the combined dataset

Table 2. Summary of tef-1 and rpb2 data sets

Datasets

Properties tef1 1st tef1 2nd tef1 3rd tef1introns rpb2 1st rpb2 2nd rpb2 3rd

Alignment size 208 209 209 404 231 231 230

Excluded characters – – – 20 – – –

Model selected GTR+I+G GTR+I+G GTR+G GTR+I+G GTR+G SYM+I HKY+I+G

Likelihood score(neg.)

799.2458 625.9714 4119.1670 5196.3535 779.8090 546.2170 4380.7173

Base frequenciesFreq. A = 0.3381 0.3065 0.1313 0.2418 0.2740 Equal 0.1910Freq. C = 0.1924 0.2315 0.3131 0.2411 0.2203 Equal 0.2647Freq. G = 0.3633 0.1914 0.2635 0.1886 0.3592 Equal 0.2274Freq. T = 0.1061 0.2705 0.2921 0.3285 0.1465 Equal 0.3168

Proportionof invariable sites 0.6988 0.6827 – 0.1355 – 0.8199 0.0597

Gamma shape 1.2044 0.5479 2.2089 4.6272 0.2463 – 3.6622

Test of substitution saturationIss 0.606 0.042 0.024 0.235 0.262 0.052 0.016Iss.cSym 1.475 1.266 1.287 1.226 0.814 0.783 0.781P (Sym) < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001Iss.cAsym 1.736 1.381 1.418 1.313 0.603 0.548 0.544P (Asym) < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001

Note: Iss: index of substitution saturation. Iss.cSym: critical value for symmetrical tree topology. Iss.cAsym: critical valuefor extremely asymetrical tree topology. P: probability that Iss is significantly different from the critical value (Iss.cSym orIss.cAsym).


Fig. 1 (a = left page; b = right page). The 50% majority-rule consensus tree from Bayesian inferenceof the combined tef1 and rpb2 sequences. Thickened branches in bold represent ML BS support greaterthan 75% and BPP greater than 0.95; thickened branches in grey denote branches supported by eitherML BS or BPP; For selected nodes ML BS support value and BPP are respectively indicated to the leftand right of slashes; The new taxa are highlighted in the shaded box; Names of African-Malagasychanterelles are in red font, Malaysian chanterelles are in lilac and North American and Europeanchanterelles are in green and blue respectively. Outgroups are in black lettering. Numbers on the rightrefer to the recognized clades according to Buyck et al. (2014).

Fig. 1a

alignment had 1179 distinct patterns with a proportion of gaps and undeterminedcharacters of 23.89%.

The topologies obtained analyzing the combined dataset are highlycongruent with published trees (Buyck et al. 2013, 2014), at least for what concerns


significantly supported branches, and the Bayesian consensus tree (Fig. 1) wasalmost identical to the optimal tree inferred under the Maximum likelihood criterion(-lnL = –18214.812299,). The two inferences mainly differ for what concerns theposition of the “Afrocantharellus” subgenus, sister to C. tabernensis/C. minorsubclade in theMLanalyses, even if not supported, and basal to the mainCantharellusclade (also not significantly supported) in the Bayesian inference, and C. afrocibarius,in a sister, well supported position to C. fistulosus in the ML analysis or sister (butnot significantly supported) to a clade comprising C. fistulosus, C. conspicuus andC. diminutivus in the Bayesian inference. Our phylogenetic inferences resolvedmainly the same, well-supported clades as previously reported (Buyck et al. 2013,2014), although the relationships between these clades remain uncertain, the deeperinternodes, in all topologies, being poorly supported whatever the analyses.

Fig. 1b


Several collections from tropical Africa cluster together in a well-supportedclade (Fig. 1b, grey box). So far, this clade remains isolated but is notably distantlyrelated to all other Cantharellus species as yet reported from Africa (Buyck et al.2000, 2012, 2013, 2014, Buyck 1994, 2014, Tibuhwa et al. 2008, 2012.) or elsewhere,and for which sequences are known (Fig. 1b). Considered individually orconcatenated, all phylogenetic inferences also revealed an internal structure withinthis clade, and concordantly and confidently resolved four well-supported, terminalsub-clades (i.e. phylogenetic species, Fig. 1b, grey box). Almost all collectionspertaining to each of these four sub-clades were initially placed under existingnames. Morphological re-examination showed combinations of morphologicalfeatures unique to and characteristic of each, thereby defining four morphotypes.The critical morphological features that differentiate them are the presence of clampconnections, the spore shapes and size, the width of the terminal elements in thepileipellis and thickness of their walls, the spacing and interveination of the gillfolds, the presence/absence of squamules on cap and/or stipe, and the colors ofbasidiomes and context. We therefore concluded that these four morphotypes/cladesrepresent four distinct undescribed species belonging to an undescribed section inCantharellus subg. Rubrinus Eyssart. & Buyck. Descriptions, comments andamendations are given in the taxonomical part.

taxonomy

Cantharellus Adans.:Fr., Systema Mycologicum 1:318 (1821)= Afrocantharellus (Eyssart. & Buyck) Tibuhwa, IMA Fungus 3:33 (2012)= Goossensia Heinem., Bull. Jard. bot. État Brux. 28: 424 (1958)

The various species are discussed in alphabetical order:

Cantharellus afrocibarius Buyck & V. Hofstetter, in Buyck et al., Fungal Diversity58(1): 286 (2013) fig. 23

= Cantharellus cibarius var. latifolius Heinem., Bull. Jard. bot. État Brux.36: 340 (1966), syn. nov.

Description: For a full description of this recently described taxon we referto Buyck et al. (2013). The material cited below entirely fits this description.

Examined collections: DEMOCRATIC REPUBLIC OF THE CONGO:Katanga Prov., near Kisangwe, Mikembo sanctuary, from arable land regeneratedmiombo with Julbernardia globiflora (Benth.) Troupin, 1194 m, S11°28’56.7”-E27°39’27.7”, 14 Jan. 2013, De Kesel leg., De Kesel 5389 (BR5020184182533);ibidem, miombo with Julbernardia paniculata (Benth.) Troupin, 1235 m,S11°28.974’-E27°40.445’, 17 Jan. 2013, De Kesel leg., De Kesel 5418(BR5020184185626); ibidem, miombo dominated by Marquesia macroura Gilg,1216 m, S11°29’3.5”-E27°40’19.3”, 23 Jan. 2013, De Kesel leg., De Kesel 5476(BR5020184191511); ibidem, under Julbernardia paniculata, 1218 m, S11°28’57.8”-E27°40’23.5”, 13 Jan. 2014, De Kesel leg., De Kesel 6000 (BR5020184195632) andDe Kesel 6002 (BR5020184196660); ibidem, miombo dominated by Julbernardiapaniculata and Brachystegia wangermeeana De Wild., 1184 m, S11°29’16.9”-E27°39’15.9”, 15 Jan. 2014, De Kesel leg., De Kesel 6011 (BR5020184198725);ibidem, miombo woodland with Julbernardia paniculata, J. globiflora, UapacapilosaHutch. andBrachystegia wangermeeana, 1195m, S11°29’16.2”-E27°39’15.3”,15 Jan. 2014, De Kesel leg., De Kesel 6014 (BR5020184199753); ibidem, under


Julbernardia globiflora, 1222 m, S11°29’14.5”-E27°39’50.5”, 15 Jan. 2014,De Kesel leg., De Kesel 6019 (BR5020184200428); ibidem, in miombo with onlyJulbernardia globiflora, 1188 m, S11°29’07.0”-E27°40’22.0”, 17 Jan. 2014,De Kesel leg., De Kesel 6033 (BR5020184204549); ibidem, miombo with Julber-nardia paniculata, 1229 m, S11°28’57.7”-E27°40’26.0”, 18 Jan. 2014, De Keselleg., De Kesel 6038 (BR5020184207632); ibidem, in mixed miombo with Brachys-tegia wangermeeana, Marquesia macroura and Julbernardia paniculata, 1221 m,S11°28’56.2”-E27°40’25.4”, 23 Jan. 2014, De Kesel leg., De Kesel 6064(BR5020184213527); ibidem, miombo with J. globiflora, along dirt road, 1079 m,S11°28’42.1”-E27°39’42.6”, 21 Jan. 2015, De Kesel leg., De Kesel 6232(BR5020184224561); ibidem, miombo woodland, 2 Feb. 2012, De Kesel & Degreefleg., Degreef 889 (BR5020169386246); ibidem, Lubemba (camp Hasson), miombowith Julbernardia globiflora and Brachystegia spiciformis Benth., 1040 m,S10°54.799’-E28°31.880’, 10 Feb. 2012, De Kesel & Degreef leg., Degreef 937(BR5020169426645); ibidem, Kipopo, miombo on clayey soil, January 1959,Schmitz-Levecq 25 (BR5020032479815, ex Cantharellus cibarius var. latifoliusHeinem).

Ecology and distribution: This species was previously considered rare as itwas only known from its type locality near Chibuli in the Copperbelt province ofZambia (Buyck et al. 2013). In Katanga, i.e. 200-300 km N-NW from the typelocality, it is fairly common and occurring in several different types of woodland. Itis mostly found in older woodland on deeper and finer soils, dominated byBrachystegia wangermeeana, B. spiciformis, Julbernardia paniculata or J. globiflora,or most often in mixed stands of the latter three tree species.

Comments: This clamped chanterelle is one of the larger species of subgenusCinnabarinus Buyck & V. Hofstetter and is placed (Fig. 1a) as sister without supportto a subclade comprising several much smaller species (the Malayan C. diminutivusCorner, and the African C. conspicuus and C. fistulosus). The caespitose habit ofC. afrocibarius is a very conspicuous, fairly constant feature of this species. Withinthe genus, this type of fruiting habit is a rare phenomenon, but in Africa a similarcaespitose habit may often be observed in other species from different subgenerasuch as Cantharellus stramineus, C. defibulatus, C. mikemboensis or C. sublaevis.However, for identification purposes, the caespitose habit should always be used incombination with other characters. Caespitose C. afrocibarius can be distinguishedfrom caespitose specimens of the above-mentioned species by the complete lack ofsquamules on both stipe and cap, combined with a lamellar, orange hymenophoregradually becoming whitish orange to whitish towards the stipe.

A comparison of available macro- and microscopical data of the typespecimen of Cantharellus cibarius var. latifolius shows that the latter taxon fullycorresponds with typical C. afrocibarius, except for the caespitose habit. Althoughwe were unable to obtain sequences from this type specimen (BR, Schmitz-Levecq25, DR Congo, Katanga, Kipopo, January 1959), we consider this taxon to representan older synonym of C. afrocibarius,

Cantharellus defibulatus (Heinem.) Eyssart. & Buyck, Documents Mycol. 31:55 (2001) Figs 2-4, 24

Mycobank: Typification MBT372679.= Cantharellus cibarius var. defibulatus Heinem., Bull. Jard. bot. État Brux.

36: 339 (1966)A full description of this ill-known taxon is given by Heinemann (1966)

and additional comments can be found in Buyck (2014, see the commentary relating


to C. sublaevis). The material cited below entirely fits the original description andwe have chosen one sequenced collection to represent the epitype, which is describedin more detail below.

Epitype: DEMOCRATIC REPUBLIC OF THE CONGO: Katanga Prov.,near Kisangwe, Mikembo sanctuary, nearly monospecific Brachystegia longifoliaBenth. miombo woodland mixed with fewMonotes katangensis (DeWild.) DeWild.,1187 m, S11°28’41.63”-E 27°39’40.26”, 27 Jan. 2013, De Kesel leg., De Kesel 5511(EPItYPUS hic designatus: De Kesel 5511, BR5020184192549).

fruit bodies very fleshy and firm, up to 120 mm diam. and more than80 mm high, often buried for another 30-50 mm in the soil, multipileate. Pileus

Figs 2-4. Cantharellus defibulatus. 2. Basidiospores (scale bar = 10 μm). 3. Basidia. 4. Hyphae from thepileipellis (all from De Kesel 5511, EPITYPE). Scale bar 3-4 = 50 μm.


smooth, convex to plane, becoming only slightly depressed, pastel yellow (3A5-6),then pale yellow to light yellow (4A3-5) or towards amber yellow (4B5-6), slightlyzonate towards the margin, sometimes with whitish areas when old and humid;margin smooth, first inrolled, remaining more or less inflexed, always irregularlyundulating and strongly lobed. Hymenophore decurrent, gill-folds normally spaced(10-12/cm), anastomosing and forking, transversally connected by conspicuousveins, uniformly colored, slightly paler and more yellow than the pileus (3A2-5),becoming light yellow (4A2-4). Stipe thick and rooting, most often splitting upwardsand/or giving rise to several fruit bodies from belowground, smooth, mat, whitish atfirst becoming yellowish white. Context whitish, yellow underneath the pileipellisand stipitipellis. taste mildly peppery. Smell fruity. Spore print not obtained.

Spores ellipsoid to subglobose, (6.8-)6.9-7.7-8.4(-8.8) × (4.5-)4.6-5.2-5.8(-5.9) μm {N = 67}, Q = (1.26-)1.3-1.47-1.64(-1.67), smooth. Basidia 58.7-72.3-85.9(-97.8) × (6-)6.1-7.5-8.9(-9.1) μm {N = 30}, narrowly clavulate to subcylindrical,mostly (4)5-spored. Subhymenium composed of narrow cells, 3-5 μm diameter.Cystidia not differentiated. Pileipellis composed of slender, straight to curvy, mostlythin-walled elements; terminal elements very narrow 32.3-54.8-77.3 × 2.9-4.8-6.7(-7.1) μm {N = 29}. Clamp connections absent.

Additional collections examined: DEMOCRATIC REPUBLIC OF THECONGO: Katanga Prov., near Kisangwe, Mikembo sanctuary, woodland with onlyBrachystegia longifolia, 1183 m, S11°28’33.8”-E27°39’42.0”, 25 Jan. 2014,De Kesel leg., De Kesel 6070 (BR5020184215583); ibidem, miombo dominated byMarquesia macroura, growing on the ground around a young M. macroura, 23 Jan.2015, De Kesel leg., De Kesel 6234 (BR5020184225599); ibidem, Lubumbashi,Jan. 1933, de Loose B13 (Holotype, BR5020032475770). TOGO: Central Prov.,Fazao, National Parc Fazao-Malfakassa, woodland dominated by Uapaca togoensisPax, on gravelly soil, 556 m, N08°43.145’-E0°46.332’, 20 Jul 2007, De Kesel leg.,De Kesel 4467 (BR5020163802667).

Ecology and distribution: So far considered rare but apparently widelydistributed and now known from Katanga province (DR Congo) and Togo (West-Africa). This chanterelle occurs in the DR Congo in woodlands dominated byBrachystegia longifolia, sometimes mixed with the dipterocarpsMarquesia macrouraand Monotes katangensis (Dipterocarpaceae). In West Africa (Togo) known onlyfrom woodlands dominated by Uapaca togoensis and equally presence of somedipterocarps, in this case Monotes kerstingii Gilg.

Comments: Buyck et al. (2013) did not provide sequenced collections ofC. defibulatus. This taxon was previously known only from the type and oneadditional specimen (Heinemann 1966). We checked the latter collection (Schmitz-Levecq 14) and found significant differences with the type. Based on the clearlysquamulose caps, with 9-12 μm wide pileal elements and excessively rare clamps,this second collection does not belong to Cantharellus defibulatus, but toC. pseudomiomboensis.

The epitype, and all other new collections presented here, fully correspondwith the holotype, although the holotype (de Loose B13) has slightly narrowerspores (6.7-)6.8-7.6-8.5(-8.4) × 4.1-4.6-5(-5.1) μm {N = 49}, hence a higher Q valueof 1.48-1.67-1.84. This confirms the spore measurements provided by Eyssartier(2001) for the holotype with a Q value of 1.4-1.73-2.0.

Cantharellus defibulatus is a large, multipileate and entirely smooth-cappedspecies. In the field it can only be confused with C. afrocibarius. However, the latterhas gills that are whitish near the stipe and yellow-orange near the margin. under


the microscope, the difference is very easily made as the latter species has clampsin all tissues.

Our phylogeny confirms the placement of C. defibulatus in subg. Rubrinussect. Heinemannianus Eyssart. & Buyck. It is placed sister with maximum support(100 ML BS, 1 BPP, fig. 1b) to the Malagasy C. sebosus Buyck & V. Hofstetter asalready suggested by Buyck et al. (2015).

Cantharellus guineensis De Kesel & Yorou sp. nov. Figs 5-7, 25Mycobank: MB 818087.Diagnosis: Differs from Cantharellus rufopunctatus by the presence of

clamp connections, a yellowish-orange context, hymenophore without muchinterveination, much less elongate spores and slightly narrower elements in thepileipellis.

Holotype: BENIN: donga prov., Bassila, Forêt Classée de Bassila, in oldforest gallery dominated by Berlinia grandiflora, alt 370 m, N8°59.805’-E1°38.689’,5 Oct. 2000, De Kesel 3005 (ADK3005, BR5020129217139).

Etymology: refers to its occurrence in Benin’s Guineo-Sudanian transitionzone.

fruit bodies solitary or gregarious, not fasciculate, fleshy, compact, 6-8 (9)cm diameter and up to 10 cm high. Pileus thick-fleshed, convex with centraldepression at first, becoming infundibuliform at maturity; surface deep yellowishorange (4A5) with a faint greenish hue (4B4-B5), soon paler yellowish (3A2) withdarker squamules, the latter fine, appressed, relatively dense in the centre andgradually sparser to absent near the margin; margin first inrolled and slightly lobed,soon inflexed, finally smooth, straight and upright, sharp and undulate.Hymenophoredeeply decurrent, gill folds widely spaced (4-6/cm margin), furcate, not or verymoderately interveined, mostly anastomosing near the stipe, easily removed fromthe underlying context, yellowish-orange (4A8-4A5), becoming concolorous withthe pileus or more vivid. Stipe (3)4-5(6) × 1.0-1.8 cm, more or less cylindrical orslightly tapering downwards, concolorous with the pileus, almost entirely beset withdarker (orange) punctiform squamules; basal mycelium whitish yellow to white.Context fleshy-fibrous, uniformly yellowish-orange (3A3-5), slightly paler in thestipe (age), deeper colored under the pileipellis, turning very slowly (hours) orangeor orange-brown when bruised. taste mild, not peppery. odour very strong, fruity.Spore print not obtained.

Spores broadly ellipsoid, 7.1-8-8.8(-9.1) × 5.1-5.7-6.3(-6.4) μm {N = 55},Q = 1.25-1.4-1.55(-1.58), hyaline, thin-walled, smooth. Basidia 53.5-66.9-80.3 ×(7.1-)7.4-8.7-9.9 μm {N = 25}, narrowly clavate to subcylindrical, mostly4 sterigmate, some 1-3 sterigmate with conspicuously enlarged sterigmata.Subhymenium indistinct, composed of interwoven elements of 2-5 μm diam.Cystidia not differentiated. Pileipellis composed of wider elements than the onesfrom the subtending context, often with oblique septa and slightly thickened walls;terminal cells 34.8-74.4-114 × 8.8-11.6-17.2(-20.1) μm {N = 18}, subcylindrical,constricted at the septum. Clamp connections large, present in all tissues.

Other collections: BENIN: donga prov., Bassila, Forêt Classée de Bassila,riparian woodland with Berlinia grandiflora, alt 370 m, N8°59.883’-E1°38.748’,03Oct.2000,A.DeKesel2981(BR5020129193860); ibidem,N8°59.874’-E1°38.728’,11Jun.2002,A.DeKesel3366 (BR5020152165698); ibidem,N8°59.685’-E1°38.606’,27Jun.2002,A.DeKesel3475(BR5020152173778); ibidem,N8°59.874’-E1°38.728’,28 Jun. 2002,A.DeKesel 3489 (BR5020152044450); ibidem,N8°59,001’-E1°38,631,7 Oct. 2002, A. De Kesel 3525 (BR5020152005062).


Ecology: So far only known from its type locality, i.e. a relatively intactriparian woodland dominated by Berlinia grandiflora (Caesalpiniaceae), withUapaca togoensis (Phyllanthaceae), Lonchocarpus sericeus (Poir.) Kunth,Pterocarpus santalinoides L’Herit. ex DC. (Fabaceae), Elaeis guineensis Jacq. andNapoleonaea vogelii Hook. & Planch. (Lecythidaceae) along the rivulet Akoka. The

Figs 5-7. Cantharellus guineensis. 5. Basidiospores (scale bar = 10 μm). 6. Basidia with mostly foursterigmata. 7. Hyphae from the pileipellis with clamps (all from De Kesel 3005, HOLOTYPE). Scalebar 6-7 = 50 μm.


herbaceous cover is poor or lacking. The species grows in the same habitat asCantharellus solidus De Kesel, Yorou & Buyck (De Kesel et al. 2011).

Comments: Our diagnosis of C. guineensis compares this species againstC. rufopunctatus because it is the most striking look-alike. The type was previouslyidentified as C. rufopunctatus in Eyi et al. (2011), a logical consequence of the factthat Heinemann (1959, 1966) placed woodland specimens with much shorter spores(Q < 1.6) and clamp connections under C. rufopunctatus. The presence of clampconnections in C. guineensis can be used to separate it from C. rufopunctatus, whichis the sister-species of C. miomboensis and belongs to a different clade (Fig. 1b).Macroscopically, C. guineensis can be distinguished from both latter species by itsstriking yellowish-orange context. From C. rufopunctatus it differs also in its gill-folds lacking the typically strong interveination.

Together with the three other here newly described species (C. mikemboensis,C. stramineus, C. pseudomiomboensis), it composed a highly supported monophyleticclade that is here described as a new section (see below).

Cantharellus mikemboensis De Kesel & Degreef sp. nov. Figs 8-10, 26Mycobank: MB 818088.Diagnosis: Differs from Cantharellus miomboensis by the presence of

clamp connections, a whitish smooth to minutely squamulose stipe, more spaced andlittle interveined gill folds, much less elongate basidiospores (Qmean = 1.54) andmuch wider terminal cells in the pileipellis (> 8 μm).

Holotype: DEMOCRATIC REPUBLIC OF THE CONGO: Katanga Prov.,near Kisangwe, Mikembo sanctuary, miombo woodland, under Uapaca pilosaHutch., S11°28.874’ - E27°40.105’, 06 Feb. 2012, De Kesel & Degreef leg., Degreef918 (JD918, BR5020169410484).

Etymology: Refers to the name of the type locality, Mikembo.fruit bodies solitary or gregarious, fleshy, compact, up to 12 cm diameter

and 10 cm high, sometimes caespitose with 2(3) pilei. Pileus thick-fleshed, convexto plano-convex then depressed to funnel-shaped on expansion; surface glabrous-tomentose, light yellow to dull yellow (3A5-3, 3B4-5) when young, then entirelysquamose, wax yellow (3AB5) to yellowish orange (4A5) and paler (3A2-3) towardsthe edge; margin lobed, briefly inrolled, soon inflexed to straight, sharp and undulate.Hymenophore deeply decurrent, pseudolamellate, not anastomosing, furcate andmoderately interveined, sometimes leaving smooth patches deeper on the stipe,concolorous with the cap (3A5-3), then becoming deeper yellow, butter yellow(4A4-6) to amber yellow (4B6), strongly demarcated from the stipe. Stipe (2)3-5 ×0.8-1.5 cm, cylindrical, wider and more massive in multipileate specimens, non-rooting, entirely smooth or only minutely squamulose, whitish or very pale yellow(3A2). Context fleshy-fibrous, whitish in the stipe, yellow marbled in the cap,yellow (3-4A6) under the pileipellis. taste mild. odour fruity. Spore print notobtained.

Spores broadly ellipsoid to ellipsoid, rarely slightly constricted, 7.3-8.2-9.1(-9.6) × (4.5-)4.6-5.3-6.1(-6.9) μm {N = 56}, Q = 1.37-1.54-1.71(-1.72), hyaline,thin-walled, smooth. Basidia slender, 52.8-63.9-75.1(-75.6) × 6.6-8.5-10.3(-10.6) μm{N = 20}, narrowly clavate to subcylindrical, with (3)4(5) sterigmata. Subhymeniumindistinct, composed of loosely interwoven cells of 3-5 μm diam. Cystidia notdifferentiated. Pileipellis composed of regular and slender, only slightly thick-walledelements with occasional clamps; terminal elements 33.1-57.2-93.1(-110.6) × 5.1-8.1-11.1 μm {N = 20}, subcylindrical often slightly constricted at the septum. Clampconnections present in all tissues.


Other collections: DEMOCRATIC REPUBLIC OF THE CONGO:KatangaProv., near Kisangwe, Mikembo sanctuary, miombo woodland dominated withJulbernardia paniculata, S11°28’57.7”-E27°40’26.0”, 1229m alt.s.m., 18 Jan. 2014,leg. A. De Kesel, De Kesel 6039 (BR5020184208660); ibidem, miombo woodlanddominated with Brachystegia spiciformis (old), Uapaca nitida Müll. Arg. andU. kirkiana Müll. Arg., S11°28’57.5”-E27°40’22.2”, 1220m alt.s.m., 23 Jan. 2014,leg. A. De Kesel, De Kesel 6065 (BR5020184214555); ibidem, miombo woodland,1200 m, 25 Jan. 2014, leg. M. Hasson, De Kesel 6073 (BR5020184217648); ibidem,miombo woodland, S11°29.088’-E27°40.571’, 1180 m, 31 Jan. 2012, leg.A. De Kesel& J. Degreef, Degreef 866 (BR5020169371099); Luiswishi, miombo woodland,close to Marquesia macroura, 1150 m, 15 Feb. 1990, leg. J. Degreef, Degreef 90-34(BR5020005058146).

Figs 8-10. Cantharellus mikemboensis. 8. Basidiospores (scale bar = 10 μm). 9. Basidia with 4-5sterigmata. 10. Narrow hyphae from the pileipellis with clamps (all from Degreef 918, HOLOTYPE).Scale bar 9-10 = 50 μm.


Ecology: Cantharellus mikemboensis is much less common thanC. miomboensis. It occurs in a number of different types of woodland with deepersoils. usually found under older growth of Marquesia macroura, Julbernardiapaniculata or Brachystegia spiciformis, most often mixed with Uapaca spp. It isimpossible to indicate a specific host.

Comments: Although the diagnosis of Cantharellus mikemboensis comparesit against features ofC.miomboensis, it also strongly resemblesC.pseudomiomboensis.

Based on its white and smooth to almost smooth stipe, C. mikemboensis iseasily separated from C. miomboensis. Based on its pale yellow to dull yellow capand almost concolorous yellowish scales it clearly differs fromC. pseudomiomboensis.In fact, the cap of the latter species is brownish to cinnamon brown at first, becomingyellowish and densely beset with light brown to cinnamon brown scales, especiallyin the center.

Based on its relatively large size, squamulose cap, yellow color and smallerbasidia, C. mikemboensis resembles species in subg. Rubrinus sect. Isabellinus(clade 2a, Buyck et al. 2013), particularly C. miomboensis. However, it differs fromall species in this subgenus by the presence of abundant clamp connections.

Young and massive fruitbodies of C. mikemboensis could be confused withthe smooth capped C. afrocibarius. Both taxa are indeed large and fleshy, with oneor several yellow caps, showing clamp connections in most tissues and slightlythick-walled elements in the pileipellis. The new species, however, becomessquamulose at maturity, has a deeper yellow and uniformly colored hymenophoreand shows wider terminal cells in the pileipellis.

Together with the three other here newly described species (C.guineensis,C. stramineus, C. pseudomiomboensis), it composed a highly supported monophyleticclade that is here described as a new section (see below).

Cantharellus miomboensis Buyck & V. Hofstetter, in Buyck et al., Fungal Diversity58(1): 291 (2013) fig. 27

A description of this recently described taxon is given in Buyck et al.(2013). The material cited hereunder entirely fits the description; the sequencedcollections cluster with the type.

Other collections:DEMOCRATIC REPUBLIC OF THE CONGO:KatangaProv., near Kisangwe, Mikembo sanctuary, regenerated miombo with Julbernardiaglobiflora, 1194 m, S11°28’56.7”-E27°39’27.7”, 14 Jan. 2013, De Kesel leg.,De Kesel 5392 (BR5020184183561); ibidem, miombo dominated by Julbernardiaglobiflora (young and old), 1202 m, S11°29’02.9”-E27°39’20.2”, 14 Jan. 2014,De Kesel leg., De Kesel 6006 (BR5020184197698); ibidem, miombo woodlanddominated by Brachystegia spiciformis (old) and Julbernardia paniculata (veryyoung), 1218 m, S11°28’57,4”-E27°40’24,7”, 19 Jan. 2014, De Kesel leg., De Kesel6044 (BR5020184209698); ibidem, miombo dominated by Julbernardia paniculata,1223 m, S11°28’59.4”-E27°40’26.3”, 23 Jan. 2014, De Kesel leg., De Kesel 6059(BR5020184212490); ibidem, miombo with Brachystegia spiciformis andJulbernardia globiflora, 1160 m, S11°29’02.1”-E27°40’28.9”, 16 Jan. 2015,De Kesel leg., De Kesel 6203 (BR5020184222505); ibidem, miombo dominated byJulbernardia globiflora and J. paniculata, 19 Jan. 2015, De Kesel leg., De Kesel6219 (BR5020184223533); ibidem, miombo dominated by Julbernardia globifloraand Brachystegia spp., 1176 m, S11°29.042’-E27°40.043’, 1 Feb. 2012, De Kesel& Degreef leg., Degreef 877 (BR5020169379170); ibidem, miombo dominated byBrachystegia boehmii Taub., B. longifolia & Uapaca pilosa, 3 Feb. 2012, De Kesel& Degreef leg., Degreef 895 (BR5020169390281); ibidem, Kipopo, miombo


woodland, 1250 m, S11°33’-E27°21’, 1 Jan. 1959, Schmitz-Levecq 39(BR5020032810243, ex. Cantharellus rufopunctatus var. ochraceus).

KENYA: Coast Prov., Malindi district, Arabuko - Sokoke, coastal miombowith Brachystegia spiciformis and Julbernardia magnistipulata (Harms) Troupin,S3°17.1’-E39°56.5’, 27Nov. 2004,DeKesel leg., DeKesel 3908 (BR5020162711427)and De Kesel 3909 (BR5020162721525); ibidem, near Shimba Hills, coastalmiombo, under Paramacrolobia sp. and Brachystegia sp., S4°07’-E39°30’, 30 Nov.2004, De Kesel leg., De Kesel 3936 (BR5020162814487); ibidem, Kwalé, Lungumaarea, Kayateleza, coastal miombo with Brachystegia sp., S4°07.966’-E39°30.655’,04 DeC. 2004, De Kesel leg., De Kesel 3995 (BR5020162782168).

ZIMBABWE: Harare Prov., Kutsaga (Tobacco Research Center), miombowoodland with Brachystegia spiciformis, S17°50’-E31°08’, 31 Jan. 1999, De Keselleg., De Kesel 2368 (BR5020112538180).

Ecology & distribution: Buyck et al. (2013) indicated that C. miomboensisis one of the most common chanterelles of the African miombo. The species isknown from Tanzania and Zambia (Buyck et al. 2013), reported under Brachystegia.Our records extend its distribution to Kenya, Zimbabwe and the Democratic Republicof the Congo. We confirm its occurrence in miombo woodland dominated byBrachystegia spiciformis. However, it is also frequently found under Julbernardiaglobiflora and J. paniculata. In spite of intensive surveying in Togo, Bénin andBurkina Faso, it is so far unrecorded from the Soudano-Guinean woodlands of WestAfrica. This may possibly be explained by the absence of suitable host genera, i.e.Julbernardia and Brachystegia (Caesalpiniaceae).

Notwithstanding the high similarity between C. miomboensis and some ofthe here newly described species, it is genetically not close and belongs in subg.Rubrinus, sect. Isabellinus Eyssart. & Buyck together with C. rufopunctatus andseveral other yellowish-brown African chanterelles lacking clamp connections(Fig. 1).

Cantharellus pseudomiomboensis De Kesel & Kasongo sp. nov. Figs 11-13, 28Mycobank: MB 818089.Diagnosis: Differs from Cantharellus miomboensis by the light cinnamon

brown color of the young cap, much larger and light brown scales on the cap, thewhitish squamulose stipe, slightly more spaced gill folds, and much less elongatebasidiospores (Q = 1.5), much wider (> 8 μm) terminal cells in the pileipellis andthe rare presence of large clamp connections.

Holotype: DEMOCRATIC REPUBLIC OF THE CONGO: Katangaprovince, near Kisangwe, Mikembo sanctuary, miombo woodland with Julbernardiaglobiflora, Brachystegia microphylla Harms and Uapaca kirkiana, S11°29,074’-E27°40,180’, 1210m alt.s.m., 8 Feb. 2012, leg. A. De Kesel, Degreef 927(BR5020169418565).

Etymology: The name epithet refers to the resemblance with Cantharellusmiomboensis.

fruit bodies solitary or fasciculate, only rarely gregarious. Pileus up to120 mm diam., fleshy and compact, first convex to plano-convex, very soon with acentral depression and somewhat lobed towards the edge; margin lobed, wavy, thick,for a long time enrolled, eventually becoming straight and sharp; surface at firstentirely light cinnamon brown (6C6-6D5) and felty-tomentose, very soon disruptingand showing a whitish to yellowish context (3A5) in between fine and flattened tosemi-erect squamulae, the centre often remaining felty-tomentose and light brown.


Figs 11-13. Cantharellus pseudomiomboensis. 11. Basidiospores (scale bar = 10 μm). 12. Basidia with4-5 sterigmata. 13. Hyphae from the pileipellis with rare clamps (all from Degreef 927, HOLOTYPE).Scale bar 12-13 = 50 μm.

Hymenophore consists of deeply decurrent gill folds of different lengths, well-spaced (3-5/cm margin), very rarely anastomosing, most often forked, usuallyinterveined, often bright yellow (3A4-5) and strongly demarcated from the whitishstipe. Stipe (2)3-5.5 × 1.0-2.5 cm, cylindrical or tapering towards the base, non-rooting, almost white or with a faint hue of yellow, entirely beset with very fine toinconspicuous squamulae, the latter pale brown, pale orange or whitish, the basalpart of the stipe often slowly bruising yellowish or orange. Context fleshy-fibrous,whitish in the stipe and most of the pileus, orange under the pileipellis, unchanging.taste mild. odour fruity. Spore print not obtained.

Spores broadly ellipsoid to ellipsoid, (6.3-)6.4-7.2-7.9(-8.3) × 4.4-4.9-5.5(-5.6) μm {N = 48}, Q = 1.29-1.46-1.63(-1.67), hyaline, thin-walled, smooth.Basidia slender, 41.8-56.5-71.2 × 6.3-8.2-10.1 μm {N = 18}, narrowly clavate tosubcylindrical, mostly (4)5 sterigmata. Subhymenium poorly defined, composed of


loosely interwoven cells of 3-5 μm diam. Cystidia not differentiated. Pileipelliscomposed of regular and slender, only slightly thick-walled elements, occasionallywith clamps; terminal cells 31.7-43.9-56 × 5.4-8.4-11.4(-11.6) μm {N = 14},subcylindrical often slightly constricted at the septum. Clamp connections large,not common, but most frequent on pileipellis elements (see ADK6198, very rare inADK6190).

Other examined collections: DEMOCRATIC REPUBLIC OF THECONGO: Katanga Prov., near Kisangwe, Mikembo sanctuary, miombo woodlandwith Julbernardia paniculata, J. globiflora and Uapaca kirkiana,S11°28’52.5”-E27°39’32.1”, 1088m alt.s.m., 13 Jan. 2015, leg. A. De Kesel,De Kesel 6190 (BR5020184219703); ibidem, on the ground, 16 Jan. 2014, leg. M.Hasson, De Kesel 6029 (BR5020184202484); ibidem, old Marquesia macrourastand, S11°29’14,0”-E27°39’52,1”, alt.s.m. 1097 m, leg. A. De Kesel, De Kesel6198 (BR5020184220440); ibidem, Kipopo, miombo forest, Jan. 1959, Schmitz-Levecq 14 (BR5020032474766).

Ecology: Miombo woodland withMarquesia macroura, but also regeneratedand denser miombo woodlands with Julbernardia paniculata, J. globiflora,Brachystegia microphylla and Uapaca kirkiana, all of which represent possiblehost trees.

Comments: Although the diagnosis of C. pseudomiomboensis compares thenew species against features of C. miomboensis, it also resembles C. mikemboensis,a much less common species. The young pileus of C. pseudomiomboensis is brownishto cinnamon brown or yellowish and then densely beset with light brown to cinnamonbrown scales, especially in the center. This feature separates it from C. mikemboensiswho has a pale yellow to dull yellow cap and almost concolorous yellowish scales.under the microscope very little differences can be observed, althoughC. mikemboensis tends to have slightly narrower terminal pileal elements and muchmore frequent clamp connections. Based on molecular data all three taxa are wellseparated from each other.

Clamp connections can be excessively rare in C. pseudomiomboensis. Thecollection Schmitz-Levecq 14, formerly placed by Heinemann (1966, pg. 340) underwhat is now called Cantharellus defibulatus, is such an example. Because of its verysquamulose pileus and terminal pileal elements measuring 9-12 μm in diameter, thiscollection actually belongs in C. pseudomiomboensis. The specimen depicted inSharp (2011, page 27, asC. rufopunctatus) probably belongs toC. pseudomiomboensis.

In the event clamp connections are not detected in C. pseudomiomboensisit may also be confused withC. tanzanicusBuyck&V. Hofstetter andC. miomboensis.However, spores of the latter two taxa are on average more elongate (Q > 1.6 andup to 2.0) while those from C. pseudomiomboensis range from Q = 1.46 (JD927) toQ = 1.55 (ADK6029).

Together with the three other here newly described species (C. mikemboensis,C. stramineus, C. guineensis), it composed a highly supported monophyletic cladethat is here described as a new section (see below).

Cantharellus rufopunctatus (Beeli) Heinem., Bull. Jard. bot. État Brux. 28: 396(1958) Figs 14-16, 30

Mycobank: Typification: MBT372680.= Lentinus rufopunctatus Beeli, Bull. SoC. R. Bot. Belg. 60: 160 (1928).= Cantharellus rufopunctatus var. ochraceus Heinem., Bull. Jard. bot. État

Brux. 28: 398 (1958).


Epitype: DEMOCRATIC REPUBLIC OF THE CONGO: oriental Prov.,Tshopo distr, Yangambi, Man-and-Biosphere reserve, rainforest dominated byBrachystegia laurentii (De Wild.) Louis ex Hoyle and a few Gilbertiodendrondewevrei (De Wild.) J.Leonard, alt 420 m, N00°49’11,5”-E24°31’20,3”, 18 Nov.2013, leg. A. De Kesel, De Kesel 5949 (EPItYPUS hic designatus: ADK5949,BR5020184194604).

fruit bodies solitary or gregarious, not connected from a common base,fleshy, compact, 6-9 cm diameter and up to 10 cm high. Pileus thick-fleshed, at firstconvex, quickly with central depression, finally infundibuliform; surface vividyellow (3A5-6) to vivid yellowish orange (4A5-6), sometimes zonate, the margin atfirst orange yellow (4B6-7), becoming paler yellowish; entirely beset with fine,

Figs 14-16. Cantharellus rufopunctatus. 14. Basidiospores (scale bar = 10 μm). 15. Basidia with 4-5sterigmata. 16. Hyphae from the pileipellis (all from De Kesel 5949, EPITYPE). Scale bar 15-16 = 50 μm.


mostly punctate squamules, denser and darker in the center (4B6), becoming finer,more spaced and paler towards the margin; margin first inrolled and slightly lobed,soon inflexed, finally straight, upright, sharp, somewhat wavy and subcrenulate.Hymenophore deeply decurrent, composed of gill folds normally spaced (8-12/cmmargin), furcate, strongly interveined, sometimes anastomosing near the stipe, paleyellowish (3A2-3A3) with a significantly brighter margin when young (3A4-3A6),becoming pale yellow (3-4A2), unchanging when bruised. Stipe (4)5-7(8) × 1.0-1.8 cm, frequently tapering downwards, upper part up to 2.5cm wide, often slightlycompressed, vivid yellow (3A5-6) to vivid yellowish orange (4A5-6), yellowishwhite towards the base, almost entirely beset with concolorous or darker (orange)punctiform squamules; basal mycelium yellowish. Context fleshy-fibrous, whitish,3A2-4A2 underneath the pileipellis and stipitipellis, unchanging when cut or injured.taste mild then becoming peppery. odour strong, similar to C. cibarius. Sporeprint not obtained.

Spores ellipsoid, (6.8-)7.0-8.0-8.9(-9.1) × (3.6-)3.7-4.2-4.7(-5.1) μm {N = 52},Q = 1.7-1.91-2.15(-2.2), hyaline, thin-walled, smooth. Basidia (40.5-)40.8-51.6-62.3× 5.7-7.5-9.3 μm {N = 24}, narrowly clavate to subcylindrical, mostly having 5sterigmata. Subhymenium indistinct, composed of interwoven elements of 2-4 μmdiam. Cystidia not differentiated. Pileipellis composed of elements wider than thosefrom the subtending context, frequently with thickened walls; terminal cells 19.6-32.7-53.2(-64.5) × 9.7-12.8-16.9(-17.6) μm {N = 21}, subcylindrical, moderately todistinctly constricted at the septum. Clamp connections absent.

Other examined collections: DEMOCRATIC REPUBLIC OF THECONGO: oriental Prov., Tshopo distr., Yangambi, Man-and-Biosphere reserve,mixed rainforest with Gilbertiodendron dewevrei and Brachystegia laurentii,alt 429 m, N0°52’5,6”-E24°27’23,5”, 13 Nov. 2013, leg. A. De Kesel, De Kesel5892 (ADK5892, BR5020184193577); Equator Prov., Binga, dry dense forest,Oct. 1934, Goossens-Fontana 992 (BR5020032807212); ibidem, Aug. 1929,Goossens-Fontana 878 (holotype of Cantharellus rufopunctatus var. ochraceusHeinem., GF878, BR5020032811257); ibidem, Djongo, rainforest withGilbertiodendron dewevrei, December 1925, Goossens-Fontana 500 (holotypeGF500, BR5020032805195);Maniema Prov., Parc National de la Lomami, ca. 3kmE-NEofKatopa, pristine dense rainforest, sandy soil, alt 450m, S002°43.5’-E25°06.4’,06 Apr. 2015, leg. Marc Sosef, Sosef 2804 (BR5020184226626).

Ecology: Only known from dense rainforests in Central Africa where it isfound almost exclusively under Gilbertiodendron dewevrei, to a lesser extent alsounder Brachystegia laurentii. Both tree species are often co-occurring andC. rufopunctatus tends to appear only in climax situations.

Comments: Cantharellus rufopunctatus is sufficiently well described (Beeli1928, Heinemann 1958) and its morphological characteristics are well documented,including drawings and a water color painting. The holotype is in good conditionbut its DNA is apparently too deteriorated for sequencing. C. rufopunctatus was firstdescribed from the Gilbertiodendron dewevrei forests of central Africa (Heinemann1958), together with a paler-colored var. ochraceus. However, in 1966 Heinemannreports both C. rufopunctatus and its variety ochraceus also from the miombowoodlands of Katanga (DR Congo), but mentions that the species is ratherpolymorphic as the specimens from Katanga differ from the type in having abundantclamps and a different morphology of spores and pileal elements. This weakimportance attributed by Heinemann to the presence/absence of clamps (evidencedalso, for ex., by his recognition of C. cibarius var. defibulatus Heinem.) led manylater researchers into confusion, not only increasing the number of citations in the


Zambezian and Guineo-sudanian ecozone (De Kesel et al. 2002, Buyck 1994, amongmany others) but also broadening the species boundaries set around the type.

So far none of the C. rufopunctatus collections previously reported fromthe miombo ecozone correspond to C. rufopunctatus and most probably the speciesdoesn’t occur there as already suggested by Buyck et al. (2013). The miombowoodlands, however, harbour a number of siblings , all of which are used for foodand local populations make no distinction between them.

We have equally studied the holotype of C. rufopunctatus var. ochraceusHeinem. (Goossens-Fontana 878, DR Congo, Binga, 29 August 1929; Bull. Jard.bot. État Brux. 28: 398, 1958) and, apart from a color difference, we could not detectany significant microscopic differences with the type variety. We therefore share theopinion of Eyssartier (2001), and consider it a mere synonym of C. rufopunctatus.Former records of C. rufopunctatus and C. rufopunctatus var. ochraceus from themiombo woodlands of Katanga (Heinemann 1966, pg. 344), i.e. collections Schmitz-Levecq 25bis and Schmitz-Levecq 39, were re-examined and belong to the herenewly described C. stramineus (see below) and to C. miomboensis (see above),respectively.

Cantharellus stramineus De Kesel sp. nov. Figs 17-19, 31Mycobank: MB 818090.Diagnosis: Differs from Cantharellus miomboensis by its overall straw

yellowish color, a paler whitish yellow hymenophore with widely spaced gill folds,the presence of clamps connections, less elongate spores and much wider terminalcells in the pileipellis.

Holotype: DEMOCRATIC REPUBLIC OF THE CONGO: Katanga Prov.,near Kisangwe, Mikembo sanctuary, monodominant miombo woodland withBrachystegia wangermeeana, S11°29’10.2’’ - E27°39’11.7’’, alt 1170 m, 21 Jan.2014, De Kesel 6051 (ADK6051, BR5020184210434).

Etymology: stramineus refers to the straw-coloured fruitbodiesfruit bodies solitary or fasciculate, fleshy, compact, up to 12 cm diameter

and 10 cm high. Pileus thick-fleshed, convex to plano-convex then moderately tostrongly depressed on expansion; surface greyish and powdery when young, verysoon uniformly dull yellow, straw yellow to greyish yellow (3B2-3, 4B2-3), withbrownish orange (5C4-6) squamules, the latter relatively dense and gradually smallertowards the margin; margin first somewhat lobed, inrolled, soon inflexed to straight,sharp and undulate, striate in older specimens. Hymenophore moderately to deeplydecurrent, pseudolamella furcate, moderately to strongly interveined, anastomosingnear the stipe, almost white at first, soon becoming yellowish white (4A2) to orangewhite (5A2), remaining relatively pale and never becoming deep orange. Stipe (4)5-8(9) × 1.5-2.4 cm, more or less cylindrical, concolorous with the pileus and entirelybeset with minute pale brownish squamules, basal part often widened and forminglateral fruit bodies, vigorous specimens rooting several centimeters below the soilsurface, basal mycelium white. Context fleshy-fibrous, white in the stipe, paleyellow (3A2-3) under the pileipellis, slowly bruising orange. taste mild. odourstrong, fruity. Spore print not obtained.

Spores broadly ellipsoid to ellipsoid, (6.9-)7-7.9-8.9(-9.1) × (4.7-)4.8-5.4-6(-6.3) μm {N = 78}, Q = (1.26-)1.3-1.47-1.64(-1.68), hyaline, thin-walled, smooth.Basidia 47.1-60.3-73.5 × 5.8-7.6-9.5 μm {N = 20}, narrowly clavate to subcylindrical,mostly 4(5) sterigmata. Subhymenium indistinct, composed of interwoven elementsof 2-5 μm diam. Cystidia not differentiated. Pileipellis composed of much wider


elements than the subtending context, with slightly thickened walls and occasionalclamp connections; terminal cells 30.9-53.4-75.8(-85) × 7.7-11.6-15.5(-22) μm{N = 14}, subcylindrical, constricted at the septum. Clamp connections present inall tissues.

Other examined collections: DEMOCRATIC REPUBLIC OF THECONGO: Katanga Prov., near Kisangwe, Mikembo sanctuary, miombo woodland

Figs 17-19. Cantharellus stramineus. 17. Basidiospores (scale bar = 10 μm). 18. Basidia clamped, withmostly 5 sterigmata. 19. Hyphae from the pileipellis with rare clamps (all from De Kesel 6051,HOLOTYPE). Scale bar 18-19 = 50 μm.


with Brachystegia boehmii, S11°28,558’ - E27°39,716’, alt. 1200 m, 01 DeC. 2012,De Kesel leg., Degreef 1035 (BR5020172909555); ibidem, miombo woodland withB. boehmii and B. longifolia, S11°28,712’ - E27°39,808’, alt. 1188 m, 11 DeC. 2012,Degreef 1098 (BR5020172958072); ibidem, Kipopo, miombo forest on brownclayey soil, Jan. 1959, Schmitz-Levecq 25bis (BR5020032809230).

Ecology: This species seems to be associated with Brachystegiawangermeeana on gravelly soils with lateritic crusts, either in monodominant standsor mixed with B. boehmii and B. longifolia.

Comments: Cantharellus stramineus shares its robust and caespitose habitwith C. afrocibarius, C. defibulatus and C. mikemboensis. Although the latter threedo not occur in exactly the same type of miombo habitat, they can be found not farfrom each other. In the field they can be easily separated from C. stramineus by theirstriking yellow colors and completely smooth stipe.

The oldest record of C. stramineus consists of a specimen (January 1959,M.C. Schmitz-Levecq 25bis) identified by Heinemann (1966, pg. 344) asC. rufopunctatus, a taxon that does not occur in the Zambezian ecozone.

In the field C. stramineus can easily be confused with C. miomboensis,especially when dealing with the off-white to yellow forms mentioned by Buycket al. (2013, pg292). Separating C. stramineus unambiguously from other large andpale yellowish squamulose chanterelles requires microscopic observation of sporeshape, presence/absence of clamps and the width of the terminal elements of thepileipellis.

Together with the three other here newly described species (C.mikemboensis,C. guineensis, C. pseudomiomboensis), it composed a highly supported monophyleticclade that is here described as a new section (see below).

Cantharellus sublaevis Buyck & Eyssart., in Buyck, Cryptog. Mycol. 35(1): 26(2014) Figs 20-22, 32

Mycobank: Typification MBT372681.A description of this recently described taxon is given in Buyck (2014). The

material cited hereunder entirely fits the description of the type.Epitype: DEMOCRATIC REPUBLIC OF THE CONGO: Katanga Prov.,

near Kisangwe, Mikembo sanctuary, under Brachystegia spiciformis (old) andseveral very young Julbernardia globiflora, 1222 m, S11°28’50.2”-E27°40’49.7”,22 Jan. 2014, De Kesel leg., De Kesel 6057 (EPItYPUS hic designatus: ADK6057,BR5020184211462).

fruit bodies mostly gregarious, fleshy, compact, 6-12 cm diameter, up to10 cm high, most often caespitose with 2(3) pilei emerging from a common stipe.Pileus convex to plano-convex soon becoming depressed to funnel-shaped; surfacenearly smooth, sometimes minutely fissurate towards the margin, light yellow, dullyellow to bright egg-yolk (3A7-6, 4AB7-6); margin lobed becoming irregularly andstrongly undulate, briefly inrolled, later inflexed to straight and sharp. Hymenophorealmost entirely smooth, becoming faintly to moderately ridged towards the margin,deeply decurrent and often leaving isolated patches on the stipe, pale yellow (4A3-4) near the margin, with a pinkish white (7A2) hue elsewhere, strongly demarcatedfrom the stipe. Stipe (2)3-6 × 0.9-1.5 cm, cylindrical or slightly compressd, moremassive in multipileate specimens, entirely smooth, pale yellow (3A3-4, 4A3-4),often with a rooting part that generates new basidiomes below the surface. Contextfleshy-fibrous, whitish in the stipe, yellowish (3-4A3) under the pileipellis andstipitipellis. taste mild. odour fruity. Spore print not obtained.


Spores very variable, ranging from broadly ellipsoid to elongate, oftensubreniform, 8.7-10.9-13.1(-14.1) × 4.6-5.4-6.1(-6.2) μm {N = 46}, Q = 1.57-2.04-2.51(-2.68), hyaline, thin-walled, smooth. Basidia slender, 53.9-65-80.8(-81.8) ×4.2-5.5-6.8(-7.2) μm {N = 32}, narrowly clavate to subcylindrical, with (2)3-4(5)

Figs 20-22. Cantharellus sublaevis 20. Basidiospores (scale bar = 10 μm). 21. Basidia. 22. Hyphae fromthe pileipellis (all from De Kesel 6057, EPITYPE). Scale bar 21-22 = 50 μm.


sterigmata in fairly equal frequencies. Subhymenium composed of slender cells of3-5 μm diam. Cystidia not differentiated. Pileipellis composed of regular andslender, sometimes slightly thick-walled elements; terminal elements 30.9-52.2-70.8(-82) × 3.7-5.3-7(-7.2) μm {N = 33}. Clamp connections absent.

Other collections:DEMOCRATIC REPUBLIC OF THE CONGO:KatangaProv., near Kisangwe, Mikembo sanctuary, miombo with old Brachystegiaspiciformis and young undergrowth of Julbernardia globiflora, 1205m, S11°29’08.8”-E27°40’35.0”, 27 Jan. 2014, De Kesel leg., De Kesel 6077 (BR5020184218676);ibidem, Lubemba (camp Hasson), miombo woodland, under Julbernardia globiflora& Brachystegia spiciformis, 1040 m, S10°54,799’-E28°31,880’, 10 Feb. 2012,De Kesel leg., De Kesel 5082 (BR5020184178703); ibidem, savannah woodland onKalahari sands, under Uapaca pilosa, U. nitida and few Isoberlinia angolensis(Benth.) Hoyle & Brenan, 11 Feb. 2012, De Kesel & Degreef leg., Degreef 942(BR5020169429677); ibidem, miombo woodland, 12 Feb. 2012, De Kesel & Degreefleg., Degreef 964 (BR5020169445837); ibidem, Fungurume; East from Koperberg,miombo woodland, 26 Feb. 1986, Schreurs1216 (BR5020008019656); ibidem,Kanonga, Upemba National Park, savannah woodland, 15 Feb. 1949, leg. Van Meel,De Witte 5542 (BR-5020032473752, ut Cantharellus cibarius Fr. var. cantharellus(Schwein.) Heim).

Ecology and distribution: This species is considered very rare and was onlyknown from its type locality near Chibuli in the Copperbelt province of Zambia andalso from one collection in the Upemba National park in Katanga, DR Congo (Buyck2014). We can confirm that it is indeed a rare species. We found it exclusively inolder woodland, mostly under old Brachystegia spiciformis, often mixed withyounger Julbernardia globiflora.

Comments: The systematic position of this species was never molecularlyestablished although Buyck (l.c.) suggested that it was probably close to theMalagasy C. sebosus and therefore belonged in subg. Rubrinus. Our phylogeny nowconfirms this placement and places this chanterelle together with other clampless,medium-sized, yellow chanterelles, such as C. defibulatus and C. sebosus, in sect.Heinemannianus.

The morphology of the epitype entirely fits the description given in Buyck(2014), although the holotype of C. sublaevis misses the base of the stipe (Buyck2014, fig. 19) and hence the protologue (Buyck l.c., pg. 26) doesn’t presentinformation on its nature. The second collection (De Witte 5542) and all of theabove mentioned collections show basidiomata being aggregated at the stipe base.Although we think the multipileate nature is a constant feature of C. sublaevis, itcannot be used as sole identification criterium since another smooth chanterelle fromtropical Africa, i.e. C. solidus, is also multipileate (De Kesel et al. 2011).

Our collections also add to the variation of spore dimensions because of themore variable number of sterigmata that we observed per basidium. Indeed, Buyck(2014) indicates predominantly 5-spored basidia [i.e. (2-4)5] in C. sublaevis. Ourobservations indicate that the frequency of 3-, 4- and 5-spored basidia is more orless equal, and that 2-spored basidia are actually not so rare. This certainly explainsthe very irregular size and shape of the spores of C. sublaevis. Buyck (2014) givesspore measurements for the type as (8.7)8.9-9.77-10.6(11) × (4.0)4.7-5.21-5.7(6.0) μm, while the range in spore size – particularly the spore length – in boththe epitype and De Witte 5542 are more important, i.e. 8.7-10.9-13.1(-14.1) × 4.6-5.4-6.1(-6.2) μm and 7.8-9.7-11.5(-14.2) × (4.3-)4.6-5.3-5.9(-6) μm {N = 49}respectively.


Fig. 23. Cantharellus afrocibarius (ADK6038, photograph A. De Kesel).

Fig. 24. Cantharellus defibulatus (ADK5511, photograph A. De Kesel).


Fig. 25. Cantharellus guineensis. Fruitbodies at different stages of development (De Kesel 3005,photograph A. De Kesel, HOLOTYPE).

Fig. 26. Cantharellus mikemboensis. Fruitbodies at different stages of development (Degreef 918,photograph A. De Kesel, HOLOTYPE).


Fig. 27. Cantharellus miomboensis (ADK6059, photograph A. De Kesel).

Fig. 28. Cantharellus pseudomiomboensis. Fruitbodies at different stages of development (Degreef 927,photograph A. De Kesel, HOLOTYPE).


Fig. 29. Cantharellus pseudomiomboensis. upper surface of a mature fruit body showing semi-erectsquamulae (De Kesel 6029, photograph A. De Kesel).

Fig. 30. Cantharellus rufopunctatus. Fruitbodies at different stages of development (De Kesel 5949,photograph A. De Kesel, EPITYPE).


Fig. 31. Cantharellus stramineus. Fruitbodies at different stages of development (De Kesel 6051,photograph A. De Kesel, HOLOTYPE).

Fig. 32. Cantharellus sublaevis (ADK6057, photograph A. De Kesel).


Cantharellus sect. Stramineus De Kesel, sect. nov.Mycobank: MB 818091.Diagnosis: Differs from subg. Rubrinus sect. Isabellinus by the presence of

clamp connections, although these are in some taxa of variable frequency andtherefore not always easy to observe.

Type species: Cantharellus stramineus De Kesel, in De Kesel et al.Cryptogamie, Mycologie 37(3), this issue, 2016, sp. nov.

Etymology: The name refers to the straw-colour of the type species.Comments: The four newly described taxa (C. guineensis, C. mikemboensis,

C. pseudomiomboensis, C. stramineus) form a significantly supported ML BS 88%,BPP 1 fig. 1b) and so far unknown monophyletic clade within subg. Rubrinus. Ourphylogeny places this new clade as sister with very high support (95% ML BS,fig. 1) to a still highly supported (ML BS 98%, BPP 1) sect. Isabellinus, and boththese sections compose the sister clade to sect. Heinemannianus with significantsupport (ML BS 75%, BPP 1). Our new section is very similar to sect. Isabellinusin being composed of medium-sized to rather large, yellowish-brown speciespossessing a squamulose cap (and stipe) surface, in contrast to species composingsect. Heinemannianus, which have a smooth cap surface and are mostly muchsmaller and also more frequently orange-red.

In contrast to both other sections of this subgenus, our new clade iscomposed of species that always possess clamp connections, although of variablefrequency and therefore not always easy to observe, e.g. in C. pseudomiomboensis.The position of this clade within subgenus Rubrinus challenges the definition ofbeing a subgenus entirely composed of species lacking clamps connections (Buycket al. 2014). As a result, the definition of the subgenus is amended below:

Cantharellus subg. Rubrinus Eyssart. & Buyck, emend. De Kesel & BuyckFruit bodies variably fleshy, large to very small; hymenophore very variable,

from nearly smooth to producing well-developed gill-folds, forking – anastomosingor not; cap and stipe smooth to strongly squamulose; hyphal endings thick- to thin-walled; clamp connections absent or present. Type: C. floridulus Heinem.

IdEntIfICAtIon KEY for CANTHARELLUSFROM MAINLAND TROPIcAL AFRIcA

Buyck et al. (2013) proposed an identification key for all tropical AfricanCantharellus. Due to the addition of 4 new taxa with clamps, clearer boundariesaround C. rufopunctatus and an increasing separation between rainforest andwoodland species, we here propose an updated/adjusted identification key fortropical African Cantharellus.

Taxa are first keyed out based on their occurrence in either woodland orrainforest. Species reported from both vegetation types have multiple entries and arefollowed by an asterisk (*) when the habitat is not the original one and their presencetherefore still needs molecular confirmation. Endemic Malagasy species are notincluded in the key.1a. Species growing in the rainforest.....................................................................301b. Species growing in woodland vegetation...........................................................2


2a. Hymenophore mouse gray and densely veined, becoming black withageing or when handled or injured. Common................C. congolensis (*)

2b. Hymenophore neither gray nor black and fruit body not blackening withage or when bruised ....................................................................................3

3a. Fruit bodies medium-sized to large, with very well developed unequal gillsthat hardly fork but may have a much lower anastomosing network in between,the whole mushroom reminding strongly of the genus Cuphophyllus(Hygrophoraceae). Clamp connections entirely absent .....................................4

3b. Fruit bodies not reminiscent of Hygrophoraceae, medium-sized to (very) small;hymenophore reticulately veined or forming well-developed, unequal, rarelyforking gill-folds .................................................................................................64a. Fruit bodies large (up to 18 cm diam.) with cap and stipe thick, firm

and fleshy, of an intense, bright orange to reddish orange or blood redthat sticks to the fingers upon handling; gills yellowish orange. Rare .........................................................................................................C. splendens

4b. Fruit bodies usually distinctly smaller and with different colors. Verycommon and widespread.............................................................................5

5a. Spores ellipsoid, Qmean = 1.6-2.0. Cap distinctly smaller and less fleshy thanin C. splendens, typically blood red and contrasting with the much more intenseyellowish gills and stipe . ................................................................C. symoensii

5b. Spores shortly ellipsoid to nearly subglobose, Qmean = 1.2-1.4. Capversicolorous, usually a mixture of gray, vinaceous, grayish red or even yellowor greenish, sometimes with clear bluish tints in the context. Gills cream-colored to pale yellow..................................................................C. platyphyllus6a. Cap and sometimes also stipe surface exhibiting, at least partly, distinct

lilac-bluish-violaceous tints; fruitbodies generally distinctly yellowingwith age .......................................................................................................7

6b. Cap and stipe without such distinct lilac-bluish-violaceous tints..............87a. Fruit body reminiscent of Craterellus, very thin-fleshed and fragile; cap

strongly squamulose; hymenophore densely veined and anastomosing, whitish.Growing in gallery forest. Rare.....................................................C. conspicuus

7b. Fruit body firm and fleshy, at least in the cap center and stipe; cap often withsome appressed squamulae; hymenophore usually not strongly anastomosingand with well-developed, unequal gill-folds, young very pale with yellowish orpinkish tints, then gradually yellowing. Rare ................C. cf. subcyanoxanthus8a. Fruit bodies pink to red...............................................................................98b. Fruit bodies without pink or red...............................................................11

9a. Clamp connections present; spores ellipsoid (Qmean = 1.2-1.4)..........C. ruber9b. Clamp connections absent; spores more variable in form, ranging from

subglobose to elongate......................................................................................1010a. Cap up to 6 cm diam.; spores elongate, Q > 2. Rare .. C. heinemannianus10b.Cap < 3 cm diam.; spores broadly ellipsoid, Q < 1.6. Very common and

widespread ..............................................................................C. addaiensis11a. Hymenophore smooth or nearly so ..................................................................1211b. Hymenophore with well-developed anastomosing veins or gill-folds ............13

12a. Clamp connections present. Basidia two-spored producing very large,ellipsoid to subglobose spores (Q = 1.1-1.4). Only known from forestgalleries in Guineo-Sudanian woodland. Rare . ..........................C. solidus


12b.Clamp connections absent. Basidia (2), 3-5 spored, spores not remarkablyvoluminous, ellipsoid (Q = 1.5-2.5). Rather uncommon ........C. sublaevis

13a. Cap smooth or matted, usually not disrupted, never squamose nor areolate .1413b. Cap disrupted, with cracks, scales, squamules or areolate..............................22

14a. Spores distinctly elongate, Qmean = 2.3, fresh fruitbodies entirely brightmandarin orange, cap 30-65 mm.........................................C. humidicolus

14b.Spores ellipsoid, Qmean < 1.8, not with this color .................................1515a. Small species, with fine stipe (< 4 mm) and cap less than 40 mm diameter ....1715b. Large species with fleshy fruitbodies, context compact and firm, with massive

stipe (> 10 mm diam), cap diameter 40-100 mm or more, usually formingcaespitose clusters .............................................................................................1616a. Cap yellow with paler, off-white areas, up to 18 cm diam. Gills whitish

near the stipe, yellow-orange near the margin. With abundant clamps ....................................................................................................C. afrocibarius

16b.Cap bright yellow orange, sometimes with off-white areas. Gills uniformlycolored, never whitish near the stipe. Without clamps .........C. defibulatus

17a. Clamps present..................................................................................................1817b. No clamps .........................................................................................................20

18a. Stipe hollow, cap 15-25 mm, yellow with brown matted centre;hymenophore with pinkish hue, terminal elements of pileipellis 3-5 μmdiameter, spore Q = 1.7. Rare...................................................C. fistulosus

18b. Stipe full, cap less than 20 mm, hymenophore without pinkish hue,terminal elements of pileipellis > 6 μm diameter, spore Q = 1.2-1.5 .....19

19a. Cap brown, 10-20 mm, gills yellow orange, terminal elements of pileipellis (8)10-15 (20) μm diam, spore Q = 1.3-1.5. Rare................................. C. schmitzii

19b. Cap yellow-orange, 8-15 mm; gills yellowish; terminal elements of pileipellis6-12 μm diameter, spore Q = 1.26. Rare .................................C. microcibarius20a. Cap vivid yellow, strongly infundibuliform, gills pale cream to vivid

yellow, Q = 1.4. Rare.................................................................. C. gracilis20b. Cap and gills not (vivid) yellow, brownish, olive brown to greyish yellow,

pileipellis tomentose; Q > 1.5........................................................................2121a. Gill folds dense, > 20/cm near cap margin, flesh greyish turning orange yellow

when bruised, pileipellis sometimes fibrillose-squamose; spores: 6-6.98-8 ×3.5-3.92-4.5 μm, Q = 1.5-1.79-2.1 ................................................C. tomentosus

21b. Gill folds not so dense, < 10/cm at cap margin, flesh whitish, pileipellisregularly tomentose; spores larger: (7)7.5-8.1-8.5(9) × (4)4.4-4.75-5 μm,Q = (1.5)1.6-1.7-1.8(1.9). Rare ..........................C. isabellinus var. parvisporus22a. Hymenophore composed of densely crowded gill folds (> 20/cm near cap

margin).......................................................................................................2322b.Hymenophore with more spaced gill folds (< 10/cm near cap margin) .24

23a. Cap brown tomentose, sometimes fibrillose-squamose; gill folds beige brownto olive brown at maturity; spores narrowly ellipsoid to elongate, Q = 1.5-1.79-2.1. Rare .........................................................................................C. tomentosus

23b. Cap distinctly squamulose-scaly, pale lemon yellow, cream to grayish or brown.Gill folds very pale, brownish to yellowish gray, remaining so at maturity,staining yellow when bruised; spores broadly ellipsoid, Q = (1.17-)1.2-1.37-1.54(-1.63). Common...............................................................C. densifolius (*)


24a. Clamps absent; spores elongate, Qmean = 1.6-2.0 ..................................2524b.Clamps present; spores broadly ellipsoid, Qmean = 1.4-1.6 .......................

................................................................................ (section Stramineus) 2725a. Fruit bodies smaller, 27-40 mm diam., thin-fleshed, fragile context; terminal

elements of pileipellis thick-walled, 5-8(10) μm diam. Possibly also associatedwith introduced eucalypts. Rare .....................................................C. tanzanicus

25b. Fruit bodies firm and fleshy, up to 12 cm diam or more ................................2626a. Stipe orange yellow and squamulose; spores elongate Q = 1.7-2.1, terminal

elements of pileipellis thin-walled and 4-7 μm diam. Very common andwidespread ...........................................................................C. miomboensis

26b.Stipe almost white or with a faint hue of yellow, entirely beset with finepale brown, pale orange or whitish squamules; less elongate sporesQ = 1.46-1.55 and wider (8-12 μm) terminal cells in the pileipellis ........................................................................................... C. pseudomiomboensis

27a. Fruitbodies straw yellowish, hymenophore pale whitish yellow. Rare.................................................................................................................C. stramineus

27b. Fruitbodies more vivid orange, hymenophore intense yellow-orange (egg yolkcolor) .................................................................................................................2828a. Context in cap and stipe entirely yellowish orange; cap and stipe surface

deep orange, densely beset with darker orange punctiform squamules;terminal elements of pileipellis (8.8-)6.1-11.6-17.2(-20.1) µm diam. Rare,under Berlinia grandiflora ......................................................C. guineensis

28b.Context white, except for a narrow yellowish orange zone beneath thepileipellis; stipe generally whitish or pale yellow, with or withoutsquamules; terminal elements of pileipellis slightly narrower: (5.1)5.6-8.3-11.2(11.3) µm diam. A complex of two morphologically similar species....29

29a. Young cap yellow, never reddish brown. Stipe whitish or very pale yellow(3a2), almost entirely smooth, sometimes with few squamules in the upperthird ...........................................................................................C. mikemboensis

29b. Young cap reddish brown, later showing yellow context between brownishsquamules. Stipe almost white or with a faint hue of yellow, entirely beset withfine pale brown, pale orange or whitish squamules ...... C. pseudomiomboensis30a. Hymenophore mouse gray and densely veined, becoming instantly black

when handled/injured or with ageing ...................................C. congolensis30b.Hymenophore neither gray nor black and fruit body not blackening with

age or when handled .................................................................................3131a. Cap and sometimes also stipe surface exhibiting, at least partly, distinct lilac-

bluish-violaceous tints; fruitbodies generally distinctly yellowing with age..3231b. Cap and stipe without such distinct lilac-bluish-violaceous tints ...................34

32a. Spores narrowly ellipsoid, 5.5-6.72-8 × 3-3.97-5 μm, Q = 1.37-1.71-2.0.Cap very dark violaceous when young; context with faint pinkish hue ..................................................................................................C. goossensiae

32b. Spores distinctly elongate, Qmean > 2 .........................................................3333a. Cap completely smooth or only wrinkled-striate or crenulate near the extreme

margin, lacking vividly yellow colors and more pinkish, lilac to cream. Sporesnarrow (ca 4 μm wide) .................................................................C. longisporus

33b. Cap radially wrinkled or striate outside the center, partly with vividly yellowcolors; spores larger ca 5 μm wide ...........................................C. cyanoxanthus


34a. Cap and/or stipe at least partly with distinct red or pink colors .............3534b.Cap and stipe lacking such colors entirely, either yellow, bright orange,

brown or gray or a mixture of these colors .............................................4035a. Clamp connections present, medium-sized species . .......................................3635b. Clamp connections absent, small to very small species..................................38

36a. Fruit bodies fleshy and firm. Cap and stipe squamulose – hirsute...........................................................................................................C. rhodophyllus

36b.Cap and stipe smooth, tomentose, or minutely squamulose....................3737a. Fruit bodies fleshy and firm; gill folds pink, becoming orange, not anastomosing,

and sometimes forked, not intervenose. Possibly conspecific withC. rhodophyllus .........................................................................C. subincarnatus

37b. Fruit bodies thin-fleshed; gill folds whitish, with yellow or pinkish hues,strongly veined-anastomosing in between .................................C. miniatescens38a. Cap not exceeding 10 mm in diameter; stipe proportionally long, at least

twice as long as cap diameter at maturity..........................C. minutissimus38b.Cap wider, 10-30(40) mm diameter; stipe proportionally shorter than cap

diameter at maturity ..................................................................................3939a. Basidia shorter than 40 μm; cap up to 40 mm diam.; spores (5.7)5.6-6.5-

7.3(8.1) × (3.4)3.5-4.0-4.5(4.7) µm.................................................C. floridulus39b. Basidia longer than 40 μm; cap up to 10 mm diam.; context reddening; spores

(6.9)7.1-7.39-7.7(7.9) × (4.0)-4.1-4.42-4.7(5.0) μm, Q = (1.4)1.5-1.68-1.8(1.9) ..........................................................................................................C. alboroseus40a. Hymenophore smooth or nearly so........................................C. cibarioides40b.Hymenophore with well-developed gill-folds or anastomosing veins ....41

41a. Fruit bodies pale yellowish ochre, ochraceous or greyish brown, ochraceousorange, never vivid yellow, orange or vivid orange........................................42

41b. Fruit bodies orange, vivid orange or vivid yellow ..........................................4442a. Hymenophore composed of crowded gill folds (L+l = 20-40/cm), originally

described from the equatorial rain forest area, often reported fromwoodlands ...............................................................................C. densifolius

42b.Hymenophore composed of distinctly more spaced gill folds. Cap andstipe with grayish yellow to brownish tones............................................43

43a. Hymenophore pale brownish with pink hue. Basidia four-spored. Spores8-8.78-10 × 5-5.82-6.5 μm. Q = 1.42-1.51-1.67...........................C. isabellinus

43b. Hymenophore safran colored. Basidia two-spored. Spores 7-8.36-10 × 5.5-6.10-7 μm, Q = 1.14-1.32-1.54 ....................................................C. croceifolius44a. Hymenophore composed of crowded gill folds (L+l = 20-40/cm) .........4544b.Hymenophore composed of distinctly more spaced gill folds ................48

45a. Clamps absent (or doubtful, see Heinemann 1958).........................................4645b. Clamps present..................................................................................................47

46a. Cap smooth, vivid orange, up to 3.5 cm diameter; gill folds 25/cm .......................................................................................................C. pseudofriesii

46b.Cap centre with punctate squamules, lemon yellow, 3-8cm diameter; gillfolds much denser (35-40/cm) .........................................C. luteopunctatus

47a. Cap up to 10 cm diam. Cap finely tomentose-squamose, margin lobate –crenulate ......................................................................................C. miniatescens

47b. Cap at most 1.5cm diam. Cap smooth, margin smooth .......................C. tenuis


48a. Fruit bodies firm and fleshy, up to 15 cm diam or more, vivid orange, withpunctate squamules all over cap and stipe surface . ........ C. rufopunctatus

48b.Fruit bodies either much smaller or not fleshy and more fragile, lackingpunctate squamules all over cap and stipe surface ..................................49

49a. Spores ellipsoid to narrowly ellipsoid, Q = (1.5)1.6-1.79-1.9(2.1); clampspresent; cap orange-yellow, often with red fibrils in cap center and extrememargin; gill folds much paler .....................................................C. miniatescens

49b. Spores subglobose, Q < 1.2; clamps absent; cap orange; gill folds concolorous ....................................................................................................C. pseudocibariusAcknowledgements. De Kesel A., Degreef J. and Kasongo B. acknowledge ASBL

MIKEMBO (Lubumbashi, RDC) and BAK (Biodiversité au Katanga, RDC) for financial andlogistic support of our fieldwork in Katanga (2012-2015). The fieldwork in Yangambi wasfinanced by BELSPO (Belgian Federal Science Policy Office) through the project COBIMFO(Congo basin integrated monitoring for forest carbon mitigation and biodiversity). MarcSosef is acknowledged for supplying material of C. rufopunctatus from the Lomami (DRCongo). The first author thanks the Leopold III-fonds voor Natuuronderzoek en Natuurbehoudvzw (Belgium) and the Stichting tot Bevordering van het Wetenschappelijk Onderzoek inAfrica (Belgium) for financially supporting the fieldwork in Bénin and Togo. Mario Amalfiacknowledges the Botanic Garden Meise for the financial support of his work. We thankCyrille Gerstmans for digitally editing the black and white plates.

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